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Proceedings of the
Linnean Society
of New South Wales
VOLUME 93
Nos. 416-418
CONTENTS OF PROCEEDINGS, VOLUME 93
PART 1 (No. 416)
(Issued 19th November, 1968)
(Presidential Address and Papers read March-April, 1968)
CONTENTS
Page
Annual General Meeting:
Report on the Affairs of the Society for the Year ae ae 1
Elections ste Be ve a nee ae we ae ee 3
Balance Sheets oF a i: a ats as ot Oa
Jounson, L. A. S. Presidential Address. -Rainbow’s Shut the quest
for an optimal taxonomy .. is : ; 05 8
Ricuarps, AotA M. The Rhaphidophoridae (Onhopicent of Australia.
Part 7. Pallidotettix, a new genus from the Nullarbor Plain, south-
western Australia... : Ee, j : ; Ne 46
SrraucHan, I. R. A taxonomic review of the genus “Mioph yes (Amira
Leptodactylidae) (Plates TAIT) 2 52
Mitton, G. W., Hineson, D. J., and GEORGE, Dy IRs (Commuinteenen by
Dr. Mervyn Griffiths.) The secretory capacity of the stomach of
the wombat (Vombatus lursutus) and the cardiogastric gland. .. 60
Hineson, Dickson J., and Mitton, .G. W. (Communicated by Dr.
Mervyn Griffiths.) The mucosa of the stomach of the wombat
(Vombatus hirsutus) with special reference to the ma
gland (Plates III-V) A as : sa £09
Kort, Parricia. A review of the genus atone Verrill, “1876 ee
RicHArDSON, LAuRENCE R. A new bdellourid-like triclad turbellarian
ectoconsortic on Murray River Chelonia He ra ee 90
CHIPPENDALE, G. M. The plants grazed by red Eaugnreon Megaleia
rufa (Desmarest), in central Australia (Plates VI-VIIT) .. so = OS
PackHAM, G. H. The Lower and Middle Palaeozoic stratigraphy and
sedimentary tectonics of the Sofala—Hill End—Euchareena region,
N.S.W. (Plates [IX-XIT) .. Re ae ee wh Ri Bras LL
GrirFIN, D. J. G., and Yatpwyn, J. ©. The constitution, distribution
and relationships of the Australian decapod Crustacea i .. 164
Davis, Gwenpa L. The embryology of EHpaltes australis Less.
(Compositae) ae ay ie ae Pale oe ar Soe eS
PART 2 (No. 417)
\ (Issued 10th March, 1969)
(Sir William Macleay Memorial Lecture and Papers read June—July, 1968)
CONTENTS
Page
Jutt, R. K. (Communicated by Mr. Rk. H. Anderson.) Aphrophyllum
(Rugosa) from Lower Carboniferous limestones near Bingara,
New South Wales. (Plate XIII) .. ae #3 he si she 193
Sranpury, P. J. Type specimens in the Macleay Museum, University of
Sydney. I. Fishes .. ea a0 ye ed of tf .. 203
DarTnaLL, A. J., Pawson, D. L., Pops, Evizaseru C., and Smit, B. J.
Replacement name for the preoccupied genus name Odinia Perrier,
1885 (Echinodermata: Asteroidea) ay a a, ci Styny Dilek
Wass, R. E., and Gouup, I. G. Permian faunas and sediments from the
South Marulan district, New South Wales. (Plates XIV-XV) 212
Moors, H. On the first occurrence of a Climacograptus bicorms with a
modified basal assemblage, in Australia A oF ae 2
MctInrosu, R. A., and Baker, E. P. Chromosome location and linkage
studies involving the Pms5 locus for powdery mildew resistance in
wheat. (Plates XVI-XVIT) 3 2 af ae aH Ae .. 232
Brarp, J. S. (Communicated by Mr. R. H. Anderson.) The vegetation
of the Boorabbin and Lake Johnston areas, Western Australia.
(Plates “ VITI- —XXIT) ne ib He a“, ie ee AZO
Fritu, H. J. Sir William J Sees a Memorial Lecture, 1968. Wildlife
Conservation ah. wo ZAC)
PART 3 (No. 418)
(Issued 18th July, 1969)
(Papers read September—November, 1968)
CONTENTS
FuuLuLEerton, R. A., and Lanepon, R. F. N. A study of some smuts of
Echinochloa spp. (Plates XXITI-XXVII1)
DarTNALL, A. J. A _ viviparous species of Patiriella (Asteroidea,
Asterinidae) from Tasmania. (Plate XXIX) .. . Si Bs
Domrow, R. The nasal mites of Queensland birds (Acari : Dermanys-
sidae, Ereynetidae, and Epidermoptidae). (Plates XXX—XXXT)
GoLpMAN, Jupy, Hiui, L., and Sransury, P. J. Type specimens in the
Macleay Museum, University of Sydney. II. Amphibians and Reptiles
Burpipen, Nancy T. (Communicated by Dr. Joyce W. Vickery.) Notes
on Vittadima triloba sens. lat. (Compositae). .. ay if ay
Kort, Patricia. A review of the family Agnesiidae Huntsman 1912; with
particular reference to Agnesia glaciata Michaelsen, 1898. ..
Sransury, P. J. Type specimens in the Macleay Museum, University of
Sydney. III. Birds. .. ae
SransBury, P. J. Type specimens in the Macleay Museum, University of
Sydney. ITV. Mammals.
ANDERSON, D. T. and Rossiter, G. T. Hatching and larval development
of Haplostomella australiensis Gotto (Copepoda, Fam. Ascidi-
colidae), a parasite of the ascidian Styela etheridgii Herdman. .
ANDERSON, D. T. and Rossirmr, G. T. Hatching and larval development of
Dissonus nudiventris Kabata (Copepoda, Fam. Dissonidae), a gill
parasite of the Port Jackson shark. ae
Abstract of Proceedings
List of Members ..
List of Plates
List of New Genera and Species
Index
SYDNEY
PRINTED AND PUBLISHED FOR THE SOCIETY BY
AUSTRALASIAN MEDICAL PUBLISHING CO. LTD.
71-79 Arundel Street, Glebe, Sydney
and
SOLD BY THE SOCIETY
1969
Page
281
294
297
427
. 439
444
457
462
464
476
482
489
497
497
. 498
ANNUAL GENERAL MEETING
27TH Marcu, 1968
The Ninety-third Annual General Meeting was held in the Society’s
Rooms, Science House, 157 Gloucester Street, Sydney, on Wednesday, 27th
March, 1968, at 7.30 p.m.
Mr. L. A. S. Johnson, President, occupied the chair.
The minutes of the Ninety-second Annual General Meeting (29th March,
1967) were read and confirmed.
REPORT ON THE AFFAIRS OF THE SOCIETY FOR THE YEAR
The Society’s Proceedings for 1967, Vol. 92, Parts 1 and 2 were published
on 11th September and 22nd December, 1967, respectively.
During the year 12 new members were admitted to the Society, three
died, three resigned and three were removed from the list of members. The
numerical strength of the Society at Ist March, 1968, was: Ordinary Members,
275; Life Members, 31; Corresponding Member, 1; total, 307.
It is regretted that the deaths of the following members have to be
reported: Miss Margery Olwyn Levy, B.Sc., Dip.Ed., died suddenly at
Katoomba, N.S.W., on 2nd April, 1967. She had been a member of the Society
since 29th April, 1964; Dr. Edward Gordon Haig Manchester, M.B., B.S.,
who was elected to membership of the Society on 29th March, 1967, died
suddenly in Sydney on 24th September, 1967; Professor Patrick Desmond
Fitzgerald Murray, M.A., D.Sc., died suddenly on a voyage to England on
18th May, 1967 (see page 3 for obituary notice); Mr. David Sutherland
North of Lindfield, N.S.W., who had been a member of the Society since 1912,
died on 20th August, 1967.
Papers read at Ordinary General Meetings totalled 27. Lecturettes were
given at the following meetings: July, Geological Aspects of the Great Barrier
Reef, by Professor W. G. H. Maxwell; October, Some QObservations on the
Status of Rainforest, by Mr. G. N. Baur. The address of the immediate Past
President, Professor R. C. Carolin, entitled “The Concept of the Inflorescence
in the Order Campanulales” was discussed at the June meeting. At the April
meeting a symposium entitled “Marine Sciences of the Central Coast of New
South Wales—Recent Researches” was held under the leadership of Miss
Elizabeth C. Pope. A symposium was held at the September meeting under
the leadership of Dr. D. T. Anderson on “Some Recent Studies on Intertidal
Animals”. Interesting notes and exhibits were given at the April, September
and November meetings. We are grateful to all who contributed in these
various ways to the interest of the meetings. No meetings were held in May
or August.
Library accessions from scientific institutions and societies on the
exchange list amounted to 1,965 compared with 2,199 and 2,016 for the years
1966 and 1965. The total number of borrowings of books and periodicals from
the library by members and institutions for the year was 222. Members and
others continued to consult publications in the Society’s rooms, and books
and periodicals were made available for photographic copying. The following
requests for exchange of publications were acceded to during the year: the
A
2 REPORT ON THE AFFAIRS OF THE SOCIETY FOR THE YEAR
Proceedings for: Folia Histochemica et Cytochemica (publication of the Polish
Histochemical and Cytochemical Society), Krakow, Poland; Annuaire, Faculte
de Biologie (University de Sofia, Sofia, Bulgaria); Atti (Societa Italiana
di Scienze Naturali e del Museo Civico di Storia Naturale di Milano, Milan,
Italy ; Botanical Reprints (instead of Proceedings) to Institute of Ecological
Botany, University of Uppsala, Uppsala, Sweden, in exchange for Acta Phyto-
geographica Suecica; Abstract of Proceedings to Science of Science Founda-
tion Library, University of Sussex, Brighton, Sussex, England. The disposal
by purchase and gift of certain periodicals and duplicates in the library has
made available considerable much-needed space, and it is proposed during the
coming year to re-arrange the library to provide adequate space for currently
received exchanges. The library itself has been greatly improved by the
installation of fluorescent lighting, a water service, sink, hot-water heater and
cupboard and a platform ladder for convenience.
The net return to the Society from Science House was $2,724.81 for the
year.
A plaster plaque of Rey. R. Collie, which has been in the possession of
the Society for a number of years, was repaired by the Art Section of the
Australian Museum and is in the Society’s rooms.
The Rules of the Society have been revised as from 29th November, 1967,
and reprinted, the date of issue being 51st January, 1968.
The Society has assisted in matters affecting the conservation of Dee
Why Lagoon, Norfolk Island and Colong Caves, and has also supported the
Australian Conservation Foundation and the Nature Conservation Council of
New South Wales.
Dr. W. J. Peacock, conjointly with Dr. D. M. Green has been awarded
the Edgeworth David Medal by the Royal Society of. New South Wales.
Dr. Peacock has been a member of the Society since 1957 and was Linnean
Macleay Fellow of the Society in Botany during 1961 and 1962.
Linnean Macleay Fellowship
In November, 1967, Miss Alison K. Dandie, B.Sc. (Hons.) was
re-appointed to a Linnean Macleay Fellowship in Botany for one year from
Ist January, 1968. Miss Dandie has continued her research on the part played
by vesicular-arbuscular mycorrhiza in crop and pasture plants in New South
Wales. Particular emphasis was placed on attempts to find a reliable means
of obtaining such mycorrhiza in test plants. Attempts to germinate Hndogone-
type spores in culture have been begun, using various methods. Hndogone-
type spores isolated from a number of different soils from Castle Hill were
compared, and found to differ only slightly in their dimensions and gross
morphology, the differences between soils being no greater than those within
soils.
Linnean Macleay Lectureship in Microbiology
Dr. Y. T. Tehan, Reader in Agricultural Microbiology and Linnean
Macleay Lecturer in Microbiology, University of Sydney, reported on his
work for the year ending 31st December, 1967, as follows: At the beginning
of this year he was in Taiwan as Leverhulme Fellow. His stay there was
chiefly engaged in lecturing to different universities. A total of over 100
lecturing hours were delivered. A number of scientific discussions and
seminars were given at several research centres. On his return to Sydney his
activities were mainly concerned with teaching and administration. However,
some research progress was made. A paper will be read in the coming
PRESIDENTIAL ADDRESS 3
Australian Conference on Electron Microscopy. Another paper on the impor-
tance of systematics of Azotobacteriaceae in the study of its ecology will be
published in the 9th International Congress of Soil Science in 1968.
PRESIDENTIAL ADDRESS
Rainbow’s End: the Quest for an Optimal Taxonomy
The aims and justification of taxonomy are discussed, followed by an exami-
nation of the foundations of ordination and classification. The Adansonian
or phenetic philosophy is critically examined and it is concluded that its
claims of objectivity and precision are ill-founded, since subjective or arbitrary
choices and definitions are necessary concerning acceptable or relevant attri-
butes, homologies and correspondences, measures and commensurabilities of
attributes, and measures of. similarity. Phylogeny, represented topologically
as a temporal branching sequence, is held to be the nearest approach to a
firm basis of reference in nature for biological classification. The charge that
phylogenetic reconstruction involves viciously circular reasoning is discussed
and rejected, though some positive feedback is admitted.
The theory and application of numerical taxonomy are discussed in
general and the potential value of numerical phyletics is stressed. The pos-
sibility of using DNA base matching as a solid foundation is briefly examined.
It is shown that, while phenetic classifications are infinitely variable, topo-
logical phyletic reconstructions do not themselves supply the kind of taxo-
nomy that is usually demanded, since they do not adequately express
significant evolutionary change of the kind expressed as “grades”. No optimal
classification can be defined, but improvement is possible up to a point of
inherent instability.
While the underlying facts and processes can be scientifically studied as
part of taxonomy, classification itself remains largely a disciplined art, which
is not convertible to an exact science by any form of arbitrary quantification.
The necessity for compromise and continued synthesis is stressed. The
suggested replacement of Linnean hierarchy by “numericlature” is mentioned
but held to be premature and to have serious disadvantages. Mathematical
concepts are discussed wherever relevant to the foundations of the subject.
(For full text see pages 8 et seq.)
No nominations of other candidates having been received, the Chairman
declared the following elections for the ensuing year to be duly made:
President: Professor T. G. Vallance, B.Sc., Ph.D.
Members of Council: R. H. Anderson, B.Sc.Agr.; Elizabeth C. Pope,
M.Se., C.M.Z.8S.; E. LeG. Troughton, C.M.Z.S., F.R.Z.S.; T. G.
Vallance, B.Sc., Ph.D.; J. M. Vincent, D.Sc.Agr., Dip.Bact.; and
G. P. Whitley, F.R.Z.S8.
Auditor: S. J. Rayment, F.C.A.
The Chairman then installed Professor T. G. Vallance as President.
A cordial vote of thanks to the retiring President was carried by
acclamation.
OBITUARY NOTICE
Patrick DESMOND FITzGERALD MurRRAY
Professor Murray, who died on 18th May, 1967, had been a member of
the Society from 1922 to 1929 and from 1949 to 1966. He was born in London
on 18th June, 1900, the son of Sir Hubert Murray, Lieutenant Governor of
Papua (1908-1940). Educated at St. Ignatius College, Sydney, he proceeded
4. OBITUARY NOTICE
to the University of Sydney, where he graduated B.Sc., with First Class
Honours in Zoology and Botany, and gained the John Coutts Scholarship
for distinction in Science, and a University Medal in 1922. Subsequent to
eraduation he spent two years working at Oxford University under Professor
Goodrich and Julian Huxley, and in conjunction with Huxley, studied the
effects of grafting certain tissues on to the membranes of the embryonic chick,
the results being given in a series of papers, three of which were published
in the Proceedings.
On returning to Sydney he was appointed to a Linnean Macleay Fellow-
ship of the Society in Zoology from 13th April, 1924, and held this Fellowship
until 1926. In 1926 he gained a Doctor of Science Degree of the University
of Sydney. After three years of demonstrating and lecturing in the Depart-
ment of Zoology at the University of Sydney he was awarded a Rockefeller
Fellowship at the Universities of Freiburg and Cambridge. In 1939 he was
appointed a London University Reader in Biology and Comparative Anatomy
at St. Bartholomew’s Hospital Medical School and Head of the Department
of Biology in the College.
In 1949 he was appointed to the Challis Chair of Zoology in the
University of Sydney, which he held until 1960, when he accepted a Reader-
ship in the Department of Zoology of the University of New England. The
two positions he held in London and Sydney were ones of considerable
academic responsibility, the duties of which he discharged conscientiously and
with distinction. Research was with him an absorbing interest and he pub-
lished many papers. Amongst other honours he was elected a Foundation
Fellow of the Australian Academy of Science.
He was interested in the affairs of the Linnean Society and was a member
of Council for five years (1950-1954). As an approachable and friendly man,
always ready to discuss matters of interest, he was held in the highest respect
by those who came to know him.
The sympathy of members of the Society are extended to his widow.
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PRESIDENTIAL ADDRESS
RAINBOW’S END: THE QUEST FOR AN OPTIMAL TAXONOMY
L. A. 8. JOHNSON
National Herbarium of New South Wales, Royal Botanic Gardens, Sydney
[Delivered 27th March, 1968]
Synopsis
The aims and justification of taxonomy are discussed, followed by an examination
of the foundations of ordination and classification. The Adansonian or phenetic philo-
sophy is critically examined and it is concluded that its claims of objectivity and
precision are ill-founded, since subjective or arbitrary choices and definitions are
necessary concerning acceptable or relevant attributes, homologies and correspondences,
measures and commensurabilities of attributes, and measures of similarity. Phylogeny,
represented topologically as a temporal branching. sequence, is held to be the nearest
approach to a firm basis of reference in nature for biological classification. The charge
that phylogenetic reconstruction involves viciously circular reasoning is discussed and
rejected, though some positive feedback is admitted.
The theory and application of numerical taxonomy are discussed in general and
the potential value of numerical phyletics is stressed. The possibility of using DNA
base matching as a solid foundation is briefly examined. It is shown that, while phenetic
classifications are infinitely variable, topological phyletic reconstructions do not them-
selves supply the kind of taxonomy that is usually demanded, since they do not
adequately express significant evolutionary changes in patterns of organisation. No
optimal classification can be defined, but improvement is possible up to a point of
inherent instability. ;
While the underlying facts and processes can be scientifically studied as part of
systematics, classification itself remains largely a disclipined art, which is not
convertible to an exact science by any form of arbitrary quantification. The necessity
for compromise and continued synthesis is stressed. The suggested replacement of the
Linnaean hierarchy by “numericlature”’ is mentioned but held to be premature and to
have serious disadvantages. Mathematical concepts are discussed wherever relevant
to the foundations of the subject.
“He said ‘I hunt for haddocks’ eyes
Among the heather bright,
And work them into waistcoat-buttons
In the silent night...”
Lewis CARROLL (1871) —
“Through the Looking-Glass, and What Alice Found There.”
The “aged, aged man” would seem to have made two misjudgements: he
sought unnecessary and rather unsuitable materials for his purpose, and then
looked in a most unlikely place for them. Is the search by some taxonomists
for the one “correct” classification of organisms equally futile and misguided ?
Before we can answer this question we shall need to look rather closely at
some of the foundations of taxonomy.
The history of taxonomy and of taxonomic attitudes and methods has
been reviewed in detail often enough. A presidential address gives one
licence to wander at large over a field of interest. That is what I propose to
do, examining certain relevant aspects or opinions, and offering comments
and animadversions upon them, in the hope of discovering or deciding what
PROCEEDINGS OF THE LINNEAN Society oF New SoutH WALES, VoL. 93, Part 1
L. A. S. JOHNSON 9
taxonomy can or should hope to achieve. I shall use the terms “taxonomy”
and “systematics” as roughly equivalent, with a tendency towards the
principles-versus-practice distinction made by some American authors (e.g.,
Simpson, 1961). Except in passing, I shall mean by taxonomy the classifica-
tion of organisms and not, as is fashionable today, of automobile components,
criminals, Latin texts (Griffith, 1967), or heraldic beasts. In order to achieve
reasonable precision, I shall have occasion at times to use some of the elemen-
tary language and concepts, but little of the notation, of certain aspects of
mathematics, in particular of set theory and higher geometry.
THe STATUS AND JUSTIFICATION OF TAXONOMY
Taxonomists are often heard to complain that other biologists accord
them insufficient respect, that universities pay little attention to taxonomy,
and so forth—in fact that they are kept in subjection in a scientific peck order
in which biology itself ranks none too high, molecular biology apart (for the
obvious reason that molecular biologists are asking simpler, and in a sense
genuinely more fundamental, questions which can be answered—sometimes—
even by physical scientists). I shall now risk the wrath of my colleagues and
assert that, although there is indeed some cause for these complaints, not
a few non-initiates pay the taxonomic priesthood more respect than we
perhaps merit. The historico-nomenclatural system, which is probably much
less justified and immutable than systematists are wont to claim, burdens
taxonomy with an enormous deadweight of out-dated and sometimes very
bad work, professional and amateur, old and new. In any other non-historical
discipline such publications would lapse into the obscurity they deserve.
Although systematics continues to attract more than its fair share of
unshakable conservatives and legalists, much of its accumulated dross must
eventually be swept away. If it is not, we may find that traditional systematics
has been by-passed, whether we like it or not. I say this as one who has
rather punctiliously followed the rules of nomenclature, though with increasing
impatience—not at the existence of rules, but at the stultifying consequences
of those we have.
Despite these criticisms, much systematic work of the past is in fact not
out-dated—just as Pythagoras’s theorem is as valid today as ever, or, less
absolutely, as Newton’s mechanics is all that we need to cope with those many
familiar problems where the scale is such that the answers given by quantum
mechanics and relativity theory would not differ, at the order of accuracy
required, from the Newtonian answers. Also, following a becalmment in the
doldrums during the first forty years of this century, systematics was able
to recruit more workers possessing sound scientific training and some idea
of the complexity and dynamics of the populations from which their specimens
are drawn. Taxonomic research improved markedly as a result.
As we all know, this revirescence was associated with the progress of
cytogenetic and evolutionary theory, and in its early days it first found full
expression in the symposium “The New Systematics” (ed. Huxley, 1940).
Much earlier, the Darwinian revolution had imbued many taxonomists
(chiefly those whose education was not completed before the establishment
of evolutionary thinking!) with the notion that if the phylogeny of a group
could be successfully reconstructed, the taxonomic problems concerning it
would be solved. Whether or not classification was thereby improved, some
life certainly came into the subject for a few decades. Other systematists
toiled on, sorting and shuffling, without much theoretical interest at all and,
when favoured by nature with clear-cut situations, some among them produced
classifications which continue to satisfy most users. There are groups in
10 THE QUEST FOR AN OPTIMAL TAXONOMY
which nobody cares except the specialists themselves. Sometimes the tech-
nicality, obscurity, and dullness of the specialists’ esoteric works have helped
to bring about and preserve this state of blessed isolation.
In general, whether good or bad, theoretically disposed or otherwise,
taxonomists themselves, like other scientists and scholars, find their own
activities self-justifying because they bring some degree of intellectual satis-
faction. This is doubtless the chief motivation for most of us, whatever
rationalizations we offer about usefulness when called upon to show why
society should support us. Nevertheless, society does support us, niggardly
though we claim this support to be. Why? Partly because, from sheer inertia,
the dispensers of funds will usually keep a going concern alive, but also,
as any worker in a systematic service organization can testify, because there
is a genuine call, from science and industry as well as many other activities
of man, for the services of the namer and classifier.
Tue NEED TO CLASSIFY
None of us can communicate, or even live, unless we sort or classify
the phenomena of our experience; further, we need in some way to label
the classes or sets to which we assign these phenomena (for an introduction
to the mathematical theory of sets see, for example, Room and Mack, 1966,
and for an application to taxonomy see Buck and Hull, 1966). Often we know
that someone else, the specialist classifier, is more efficient at this task than
we are, and so we avail ourselves of his services. Having classified and named
(labeled) a phenomenon or group of phenomena, we can ascertain further
information about it and can talk about it. We may even, naively, consider
that we know what it is. How often an inquirer asks, “What is this
organism?” and, on being told “IMelania alba’, considers himself somehow
wiser than before. Anthropologists know well the supposed power conferred
by knowledge of a name. |
The role of taxonomy in education is emphasized by Davis and Heywood
(1963), who say that it “remains the principal agency through which the
student may gain acquaintance with the diversity of organisms, the patterns
of variation in the living world and .. . the evolutionary mechanisms which
have brought these patterns about. The taxonomic approach is a focal point
for students of biology and enables them to fit together into a framework
of ideas a mass of otherwise unco-ordinated facts”.
The classificatory nature of the whole of science is sometimes stressed.
Many of my generation will remember from their schooldays a textbook of
chemistry by Sherwood Taylor (1939), which begins with the words: “Natural
Science is the process of systematically arranging and classifying man’s
knowledge of the world about him.’ Perhaps this only partially defines the
nature of science, but the search for more inclusive and if possible more
simply and generally describable classes, of relations as well as things, is an
important factor in arriving at the general but precise “laws” (as they are
so inappropriately called) which science seeks.
THE QUESTION OF OPTIMALITY
So much for the need for some sort of taxonomy. A chain of questions
then arises: Are some classifications better than others? If so, when does
the improvement (assuming that we have some means of assessing it) justify
changing an existing system—or, how are we to choose between co-existing
systems? If improvement is possible, then is there, logically, a best classifica-
tion (practically attainable or not) to which we can try to approximate?
Finally, if there is an optimal classification, what is its basis?
=F L. A. S. JOHNSON 11
As soon as we attempt to answer these questions, or even to define the
terms used in them, we find ourselves faced with many more questions. In
any field where definitions are required we are led into an infinite regress;
so we must agree to take certain things for granted—to set up starting points,
as it were. In the more rigid contexts of logic or mathematics one starts
“with certain postulates, but even in the expression of these one must perforce
use certain terms and syntactic relationships of undefined meaning. For
example, in modern mathematics, the concept of “point” is usually left
undefined; likewise, logical links such as “it follows that” must be accepted
“more or less intuitively (Courant and Robbins, 1961).
Much has been written on the philosophical bases of classification,
including biological classification. While this literature is academically
interesting and has important implications for essentially simple (but not
always easy!) domains of inquiry such as mathematics, its application to
essentially vague (i.e., very highly complex) domains such as _ biological
taxonomy seems to be of little practical assistance. Most practising systema-
tists continue their studies without worrying about philosophical foundations
and it appears that, in their own work, many philosophically minded authors
proceed in much the same way as other competent systematists. In contrast,
evolutionary theory has noticeably influenced the methods, approach, results,
and particularly the interpretations, of those systematic workers who are
strongly aware of the genetical and evolutionary background and implications
of their work, whether they call themselves “biosystematists” or eschew such
labels.
Tur PHILOSOPHY OF PHENETICISM
Despite this lack of practical effect, the modern philosophical school
of taxonomy, foremost among whose early spokesmen were the botanist J. S. L.
Gilmour (e.g., Gilmour, 1940), now at Cambridge, and (somewhat later) the
Oxford zoologist A. J. Cain (e.g., Cain and Harrison, 1958; Cain, 1959),
has had an increasing appeal over the past thirty years to theoretically
minded taxonomists (especially those educated in certain universities). The
background to this way of thought is the “operational” approach of logical
positivism, a more far-reaching anti-metaphysical philosophy than empiricism
but, like empiricism, of obvious appeal to the scientific mind (Britton, 1958,
and references therein and in Gilmour and Walters, 1964, and Carolin, 1967) .*
It has culminated in the so-called Adansonian, neo-Adansonian, or phenetic
credo which, perhaps by historical accident, is now so closely associated with
what is labelled numerical taxonomy (Sneath and Sokal, 1962) or taximetrics
(Rogers, 1963). The strong influence of the phenetic viewpoint is exemplified
- * To define my own philosophical standpoint, I should perhaps say that, while
preferring logical positivism to metaphysical philosophies, I find it entirely reasonable
to believe in the material reality of the universe—material, of course, in the sense
that, despite the Uncertainty Principle, we can learn a great deal about the behaviour
of “matter-energy” at various levels of aggregation and organization, and that the
fundamentals (though not always the details) of this behaviour are independent of the
existence of ourselves or other observers. As to the place of purpose and particularly
of life in this physical world, I can see no justification for any form of supernaturalism,
mysticism, transcendentalism, or neo-vitalism, nor can I see anything inconsistent with
physics in the fact that, in the sense of information theory but not of physical thermo-
dynamics, organisms increase negentropy while they grow as individuals or populations.
Life is the state of possessing mechanisms for self-perpetuation, replication, and (in a
sense) occasional increase of the organization of systems of physical components; that
is the only way in which it is “different”. Explicitly or not, these views are probably
shared by most biologists. Neo-vitalism today seems often to arise, curiously enough,
among physicists (e.g., Wigner, reviewed by Pais, 1967).
12 THE QUEST FOR AN OPTIMAL TAXONOMY
in the generally well-balanced and comprehensive modern textbook of angio-
sperm taxonomy by Davis and Heywood (1963), where its merits are accepted
as almost self-evident.
Strictly, Adansonianism advocates multi-attribute classification, grouping
on the basis of many equally weighted attributes drawn from many parts of
the organism, and refusing to attach greater weight a priori to certain
“essential” characters. Michel Adanson’s approach (Lawrence, 1963), in the
context of the eighteenth-century Enlightenment, represented a deliberate
break from the blend of debased Platono-Aristotelian philosophy and theology
which constituted mediaeval Scholasticism. Although it was abandoned by
progressive philosophers, Scholasticism continued to obfuscate some scientists’
thought long after the Middle Ages and its outward forms, at least, were
preserved in the concepts of “genera”, “species”, “characters”, and “dif-
ferentiae”’, as well as in the doctrine of divine special creation, accepted by
Linnaeus and other orthodox taxonomists of his day (see also. Hull, 1965).
In practice, many good systematists were partial Adansonians long before
the term came into its present vogue. Phylogeny, as we understand it, was
a concept unknown to Adanson, though he was no divine-creationist, but
modern pheneticism not only purports to be stringently Adansonian but also
emphatically rejects the use of phylogenetic considerations in reaching
taxonomic conclusions (that is to say, in arriving at particular classifications).
However, its proponents are mostly careful to state that they do not reject
subsequent or independent phylogenetic interpretation, or “speculation” as
many prefer to put it. Most pheneticists also reject taxonomic use of the
“biological species concept” developed, with variations, in numerous publica-
tions by Dobzhansky, Huxley, Mayr, Stebbins, and others, and strongly
advocated in rather purist terms by Askell Live (e.g., Dobzhansky, 1951 ;
Huxley, 1942; Mayr, 1939, 1957a, 19576, 1963; Stebbins, 1950; Love, 1964).
Since many evolutionary taxonomists and biosystematists have thought
that pheneticism unjustifiably neglects the important gains in understanding
contributed by their approach to biology, a spate of discussions has ensued,
some of them strongly polemical, in which various issues have been debated
and frequently confused. Traditional taxonomists have joined in with more
pragmatic arguments and from fear of displacement or interference by tech-
nical innovations which, perhaps often rightly, they regard as unnecessary.
On the whole the pheneticists have spoken more loudly and with the confidence
of revolutionaries who sense that the Zeitgeist is on their side. In the
course of a decade the numerical pheneticists have come to hold the centre
of the stage. Such successes are at times due to salesmanship rather than
scientific merit, and we shall do well to look critically at the foundations
of phenetics. Time and performance, of course, will provide the acid tests.
I shall defer most of the discussion of numerical techniques in taxonomy
until we have dealt with the more general subject of systematizing the objects
of our experience.
Tue NATURE OF ORDINATION
As several authors (e.g., Williams, 1967) have recently reminded us, an
orderly arrangement of objects, or more usually of the symbols representing
them, need not be a classification; we may settle for an ordination, that is,
we may assign relative positions to objects according to their states with
respect to a set of their “attributes”, a separate dimension being MESES
for each attribute. This procedure establishes an “attribute-space” (usually
abstract); in which the objects are represented by points or sometimes
regions. Obviously, to be meaningful in the ordination, each of these
= L. A. S. JOHNSON 13
attributes must be represented by at least two states in the set of objects
under consideration; in other words, they are “pluri-state attributes”.* An
example of the simplest non-trivial case would be the ordering in one
dimension of, say, men according to their measurements (states) of the
attribute height, or of events according to sequence in time (unless relativistic
‘considerations are negligible the latter case requires specification of a frame
of reference). Two or more objects may occupy the same position in the
attribute-space. They are then indistinguishable with respect to the attributes
concerned, but we may still regard them as distinct entities by reference to
- other attributes. ,
Ordination does involve some implicit classification, as we shall see.
Consider a set of objects each possessing what we may call “elementary
attributes”. For the moment we may define the latter as any properties we
can and wish to specify, for example six-leggedness, a length of four milli-
metres, or orange colour. Before we can assign positions to the objects in
any ordination we must establish working homologies between them. This
implies setting up one-to-one (pluri-unique) correspondences, over the
object set, between some of the elementary attributes of the objects. The
result may be more or less reasonable according to the circumstances.
It is reasonable to establish a correspondence between the surface areas of a
moss leaf, a cycad leaf, and a lycopod leaf if we are interested in photo-
synthetic capacity—it is scarcely reasonable if our interest is in phylogeny,
since most botanists regard these leaves as evolutionarily non-homologous.
The establishment of a correspondence ranging over the object set, whether
reasonable or not, is tantamount to a classification of the attributes: jwe have
assigned a certain one of the elementary attributes of each object to a class.
From the property by which we define this class we derive a pluri-state
attribute applicable to all members of the object set; the elementary attributes
then become the individual “states’’.
The property “measurable length” is here regarded as an abstraction from
the set of actual measurable lengths; the very definition of this set implies
a classification of these elementary attributes on the basis of a property
common to them all (the property of being a length). We could look at this
relation in other ways: for instance, we might regard the possession of a
particular length as dependent on having length at all; again, the particular
lengths are attributes of the individual objects while length is a property
of all the objects.; It is unfortunate that the term “attribute” (or
“character”) is used in the literature at these different levels. In what
follows it will be necessary to use it in both senses but either the context or
some qualifying term should make the particular application clear.
It has been assumed in the preceding that for a single object the “states”
should be mutually exclusive—ordinarily a leaf cannot have more than one
measurement of length (at any instant). When the actual observations are
made upon parts of the objects under ordination (or classification) the state
may be expressed by some suitable statistic, for example, the mean leaf-
length or the largest observed leaf-length (the object here being a plant, a
*T shall use the prefixes “pluri-”’ and ‘multi-’ to signify respectively “more than
and “more than two” (or “more than three” for dimensionality).
+ When appropriate, the zero state must be included in the range of the pluri-state
attribute. Here length, even if sometimes zero, is considered to be a property of all
the objects if a comparison on the basis of length is meaningful. Thus, for a snake,
it is reasonable to treat the length (zero) of the external limbs as a comparable
attribute to the non-zero lengths of such organs in other reptiles; on the other hand
for an amphioxus, say, the concept of a measure of length, zero or not, of limbs
is inapplicable. In the latter case, if comparing with other chordates, the two-state
attribute “presence or absence of limbs” might be appropriate.
”
one
14 THE QUEST FOR AN OPTIMAL TAXONOMY
population, or a leaf over an extended time). Although this procedure will
simplify the ordination, it will result in loss of information. Alternatively,
the objects (being composite with respect to the attribute or attributes
concerned) may be represented in the attribute-space not as single points but
as regions, which may overlap. For complete specification we then require
an evaluable density function. For any one attribute (dimension) this may
or may not be easily expressed. If the distribution of variation is Normal,
we need to specify only (estimates of) the mean and the standard deviation
(or the variance) ; if it is non-Normal but follows some other regular pattern
it may also be expressed in terms of the distribution function and a small
number of parameters. Sometimes a logarithmic or other non-linear scaling
transformation may yield such a simple distribution. However, frequency
(=density) distributions of variation may be quite irregular, or plurimodal,
or discontinuous (e.g., flowers 4-merous 21%, 5-merous 79%). Moreover,
since more than one attribute is usually involved, the covariation within the
composite object also becomes important. Thus, the pluridimensional density
functions may be very complex indeed, and the extent to which they should
be simplified for composite objects (e.g., species or higher taxa) will be a
matter of judgement and practicability in each case.
Serious difficulties are introduced into the dimensional representation
if some (pluri-state) attributes depend. for their expression on the existence
of certain states of others, and are therefore not uniformly relevant over the
whole set of objects. These circumstances result in the attribute-space being
inhomogeneous as to dimensionality (see also Reynolds, 1965). In numerical
taxonomy this is a practical problem which has usually been somewhat
unsatisfactorily avoided by various shifts and devices such as redefinition of
attributes, or by weighting methods (Kendrick, 1965) which most numerical
pheneticists frown upon (see Long, 1966, for a facile dismissal and misunder-
standing of the serious dependence problem raised by Kendrick).
Still other problems arise when “states” which it seems reasonable to
group into a single “attribute” do not admit of an unequivocal measure by
which they may be arranged serially, for example, variants of a chemical
constituent which differ in replacement-groups. A representation which pre-
served the symmetry of such cases, and in which they could be combined with
serial (or two-state) cases would call for a space wherein some attributes
were expressible in terms of linear co-ordinates and some in terms of higher-
order symmetrical (e.g., triangular, tetrahedral, etc.) co-ordinates. This of
course implies dimensions within dimensions and enters realms of complexity
which, though perhaps beyond practical handling and certainly introducing
anisotropic properties into the spaces concerned, should not be glibly passed
over, since they are inherent in the general problem of ordination. Anticipating
matters to be discussed later, in numerical classification, which is not neces-
sarily subject to the same practical restrictions as to the spaces implied in
the models used, Lance and Williams (1967c) have devised an information-
statistic computer strategy to deal with mixed data including non-exclusive,
non-serial,* multi-state attributes. Still more recently, Wallace and Boulton
(1968) have developed another mixed-data strategy.
Although ordination implies assignment of relative position, it is not
essential for it to be metric; no particular fixed measure is obligatory. The
relation expressed by saying that, in a one-dimensional linear (in the sense
of non-closed) space, B les between A and C does not require either that we
* “Non-serial” seems preferable in this context to “disordered”, as used by Lance
and Williams.
5 L. A. §. JOHNSON 15
know just how far B is distant from A and C, or that it has any fixed
position, or indeed that a measure of distance has any meaning at all. Here
three distinct points will define an ordering, without any specification of
direction in the space. If a direction is assigned, two points A and B are
sufficient to define an ordination. Further, ordination remains mathematically
meaningful, though difficult to handle, if (in each or any of the dimensions)
the arrangement is cyclic (then, however, we should require a minimum of
four points in the non-directional non-metric case, three in the directional
non-metric) or in some other way not simply linear. Moreover, no concept
of a continuum is necessarily required; there may be only discrete positions.
Thus, the abstract space in which we represent an ordination may differ from
our everyday concept of physical space in one or more of five ways:
(1) it may be finite,
(2) it may be more than three-dimensional,
(3) it may be non-Euclidean,
(4) it may be non-metric,*
(5) it may be discontinuous.
It must, however, be a topological space. In loose terms: however we may
deform our representation by variation of scales in any direction, the ordina-
tion must not be altered. In any but the simplest cases it is no easy matter
to elucidate and explain the topological and metrical properties of the infinite
variety of definable spaces; this is no reason to assume, as is often done,
that simple spatial models are particularly appropriate to taxonomic
ordination. (For an introduction to some abstract spaces see, e.g., Sawyer,
1955, and Courant and Robbins, 1961; there are many more advanced texts,
for instance that of Kelley cited by Williams and Dale, 1965).
Non-mathematicians are conditioned to believe that there is something
‘natural’ about Euclidean space and the Euclidean metric (the latter means,
roughly, that if we set up rectangular, similarly-scaled, Cartesian co-ordinates
in an n-dimensional space then the distance between any two points P and Y
is given by the Pythagorean function
AUPQ=LE (ip @0))
where %;p, %,9 are the co-ordinates of P, Q respectively in the ith dimension.
This is an extension to n dimensions of the familiar theorem of Pythagoras
that the length of the hypotenuse of a right triangle is the square root of
the sum of the squared lengths of the two orthogonal sides. (Oblique
co-ordinate axes may also be set up in a Euclidean space, and are used with
the Mahalanobis “generalized distance” measure (Sokal, 1965; Menitskii,
1966) though Kendall (1957) and Williams and Dale (1965) have challenged
the validity of the procedure in respect of this particular statistic.) However,
from what we may know of cartography and navigation most of us will
admit that something rather different is involved in the geometry of the
surface of a sphere. We customarily think of a sphere as an object in three-
dimensional linear (Euclidean) space. Mathematically, however, its surface
geometry can be considered in complete isolation as a closed, curved, two-
dimensional space which we can, if we wish, embed in a three-dimensional
Euclidean space. We shall probably also have heard that representation of
space-time in the General Theory of Relativity calls for a particular four-
dimensional case of a strange collection called Riemannian spaces, which
* Conditions exist in which there is not a complete absence of metrical properties:
semi-metric, quasi-metric, and disjoint metric spaces all arise in numerical taxonomy
(Williams and Dale, 1965).
16 THE QUEST FOR AN OPTIMAL TAXONOMY
constitute a more general class of curved spaces. These notions of naturalness
arise in the ordering of our spatial experience of the physical world, but we
are accustomed also to graphical representation of non-spatial quantities in
co-ordinate systems in what appear to be Euclidean spaces. Actually, such
representations often do not imply, or allow, any concept of inter-point
distance involving more than one co-ordinate, and there is then in fact no
implication of a Euclidean (Pythagorean) metric. As an extension from
such representations, the concept of abstract Euclidean spaces of more than
three dimensions has become familiar, as has the correspondence (iso-
morphism) of algebra and geometry, first clarified by Descartes, which freed
geometry of the need for pictorial representability.
Euclidean metrics are implicit in most of the models used in classical
statistics, including such probabilistic techniques of multivariate statistics
as principal component analysis, factor analysis, and canonical analysis, which
depend on the method of least squares (e.g., Kendall, 1950, 1957; Seal, 1964).
Because of these familiarities and the convenient pre-existence of statistical
techniques, Some authors on taximetrics (e.g., Sokal, 1961, 1965; Sokal and
Sneath, 1963; Goodall, 1964; Jancey, 1966) have shown a strong preference
for Euclidean models. Sokal (1961, p. 73) and Boyce (1964) speak (nonsensi-
cally in the context of comparison of attribute sets) of the Pythagorean
measure as “true distance” when contrasting it with the “mean character
distance” (M.C.D.), used by Cain and Harrison (1958). The M.C.D. in fact
defines what may be called a lattice metric—the distance function here is
EI) | Zip—®i9 |
; : : 1 ,
(ignoring the M.C.D.’s sealing factor ‘| which means that the shortest
distance between two points is, in general, “around the corners”, as for a
rook’s moves in chess (#%=2) or an ant travelling along the bars of a
children’s “jungle-gym” (n=3). Such a lattice-metric space can be embedded
in a Kuclidean space; it differs from the latter in that, given a basis of
rational numbers, no concept of algebraic irrational numbers is needed to
express distances within it, nor are there any smooth curves or transcendental
irrationals like 7. Unlike those of a Euclidean space, lattice-space distances
are not invariant under rotation of axes, indeed in general (with a rational-
number basis) the “fixed” points themselves will ‘disappear’? under such
rotation. These details are not relevant to practical taximetrics but are
mentioned to illustrate that some of the mathematical properties which we
take for granted in the Euclidean metric can change drastically with a very
simple change of metric. Lance and Williams (1967c) refer to this lattice
metric as the “Manhattan metric” and point out that it is the first-order case
of a general class (Minkowski metrics), the Euclidean metric being the second-
order case.
The infinite-order case is of interest since it is the “chess-king’s
metric” in which
ad(P,Q)=| Uip— Vig | sae.
that is, the distance is simply the value of the greatest single co-ordinate
difference between the points.
There is no @ priori reason why such a metric, or indeed many others,
should yield a less “realistic” measure of “distance” (or its complement,
similarity) between sets of attribute-states. Nevertheless, even the mathe-
matically sophisticated Williams and Dale (1965) favoured Euclidean
systems, on grounds of statistical convenience and “pictorial” representability
(which fails in any case due to the distortions and indeed the semi-metric
= L. A. S. JOHNSON ily¢
property of projective mappings of higher-dimensional spaces on to their
lower-dimensional subspaces, i.e., originally distinct points may not be distinct
in the mapping). Jancey (1965) suggests that the intuitive concept of simi-
larity in terms of real spatial relationships renders the Euclidean metric
worthy of retention. If this merely reinforces preconceptions, its value seems
doubtful and, as Macnaughton-Smith (1965) says, “this ‘visualizable’ quality
is by no means a necessity, and one would wish to use the most appropriate
function regardless of whether it was visualizable”. More recent papers on
taximetrics (e.g., Lance and Williams, 1967c) show less attachment to
Euclidean representations.
For particular purposes, ordination procedures may be more efficient
than classifications in that they need not involve us in loss of information,
nor do they set up so many “artificial” distinctions or segmentations of
gradual transitions. Their usefulness will naturally depend on the appro-
priateness to our purposes of the attributes chosen, and on the measure
adopted for assigning position in each dimension. Incommensurability in the
various dimensions is a problem only if we are interested in such concepts
as Similarity over a multiplicity of attributes or “distance” between objects
(points or-regions) in the attribute-space. Perhaps, if our mental apparatus
were differently organized, multidimensional ordination would meet most
of our requirements for the organization, retrieval and comparison of
information. This is a question for those studying the design of logical and
quasi-mental machines.
Tuer NATURE OF CLASSIFICATION
As it happens, our minds cannot cope with multidimensional systems
and we need to classify, even though we lose information and introduce
distortions and artificialities in the process.
In dealing with the chain of questions posed earlier, it will be useful
to refer frequently to one of the most definitive statements of the “philo-
sophical” attitude, a contribution by Gilmour and Walters (1964) entitled
“Philosophy and Classification”. I should make it clear that, although I
shall offer some severe criticisms of the purely phenetic approach, a good
deal of what Gilmour and Walters have to say is not in dispute and that
they are aware of some of the unresolvable aspects of the problems raised.
It has been repeatedly stressed, especially by Gilmour and his followers,
that the values of classifications should be assessed according to the range
of their purposes. We need not here discuss classifications of narrowly-
defined purpose, although many of the problems associated with more general
classifications may arise, in parvo, in the special cases also.
Biological taxonomy is expected to produce classifications of broad
utility; Gilmour and Walters have termed these general-purpose classifica-
tions. More essentialistic taxonomists have aimed to produce _ so-called
“natural” classifications which, it is claimed, would in part meet the require-
ments of a general-purpose classification but would also, as it were, reflect
some more fundamental truth about nature. As Gilmour and Walters point
out, there are philosophical objections to the term “natural” (and its opposite,
“artificial”) in this usage. They say: “the view that there are such ‘natural
kinds’, differing in some ‘fundamental’ way from other, ‘artificial’ methods
of classifying the same objects, is very difficult to sustain; a more useful
way of looking at the situation is that these so-called ‘[natural] kinds’ are
classes showing [a] high degree of correlation of attributes, differing only
in degree from other classes with a less high correlation”. (The second part
of this statement raises difficulties, discussion of which is deferred.) Later,
18 THE QUEST FOR AN OPTIMAL TAXONOMY
they outline eight principles which they suggest should be applied to bio-
logical classification. Since these represent, in summary, an influential
attitude, I shall quote them in full as a framework for critical comment.
“(1) The term ‘classification’ is used by philosophers to describe the
act, conscious or unconscious, of grouping objects into classes because
‘of certain attributes they have in common.”
Comment: We may have difficulty in defining “objects” and “attributes”
but the definition will serve. Here, and in what follows, these authors use
“attributes” to include what I have called “elementary attributes” rather
than “pluri-state attributes”. I shall follow their usage while it remains
relevant.
“(2) Classification, used in this sense, is man’s basic method of
dealing with the multiplicity of individual objects in the world around
him.”
Comment: Whilst the statement is true enough, its restriction to “man”
is unnecessary and symptomatic of an anthropocentric viewpoint which
pervades philosophy. Other animals classify, so do some machines; certainly
any highly intelligent being would do so (though, as I have said, some might
be less compulsive classifiers than ourselves). The process of classification
does not depend on the existence of man. It does, I suppose, depend on the
existence of an agent of some kind—“classifico ergo sum!’—but we may be
led into unprofitable by-ways if we pursue this further.
“(3) Since classification is a product of man’s need to deal with
his environment, the actual classifications that he makes are determined
by his desires and purposes in relation to that environment.”
“(4) The suitability of any particular classification can only be
judged in relation to the purpose for which that classification was made.”
Comment: This is not necessarily so; a classification made for one purpose
may suit another very well. It should be judged in relation to the purpose
for which it is required.
“(5) Two types of classification can be distinguished, with every
gradation between them: ‘general-purpose’ classifications and ‘special-
purpose’ classifications.”
“(6) General-purpose classifications consist of classes containing
objects with a large number of attributes in common, thus making them
useful for a wide range of purposes; special-purpose classifications
consist of classes containing objects with only a few attributes in common,
and hence serve a more limited range of purposes.”
Comment: In a “common-sense” way, we doubtless all think that we
can see what is meant here. However, the concept of “number of attributes
in common”, or more sophisticated measures of similarity, lies at the heart
of the phenetic (Adansonian) approach, and of the mathematical methods
which have been developed on the basis of this approach. Therefore we must
examine this statement more critically. There are several hidden variables
here, and unless we can somehow define, measure, and control them, the
statement simply will not do as a basis for the precise, “objective” approach
to taxonomy which many pheneticists state to be their aim.
First, we must consider the domain from which the attributes are to be
chosen. Every “object”, a term which may be extended to cover any physical
or conceptual object of discourse or thought, has an infinitude of attributes.
That is to say, the object itself does not set any bound beyond which we
can say “no further attributes exist”; this conclusion is not affected by the
L. A. §. JOHNSON 19
practical limitations of our thought. (We can ignore such playthings of
paradox-fanciers as “the concept which has no attributes”.) A little reflection
will show, for instance, that any object bears various relationships, tenuous
as they may be, to every other physical object or collection of objects in the
universe, at every point on the world-line of every particle in space-time.
There are likewise relationships to the past and future states of the object
itself, and indeed to innumerable abstract concepts. Perhaps this sounds
extreme, but it is an inescapable conclusion from any general concept of an
attribute, and there is no @ priori reason to stop at any particular point in
our search for further attributes. It may be objected that relations to other
objects are not intrinsic attributes of the object under consideration.
Reflection will show that all describable characteristics are relations to other
objects or concepts; we simply cannot speak meaningfully about the pro-
perties of a thing in itself, Plato or Whitehead notwithstanding. Likewise,
it is of no practical help to adopt a holistic standpoint and claim that every-
thing is part of one great integrated whole and that the individual objects
and their attributes are mental abstractions from this whole—this can only
lead one to say “everything is as it is’ and to resign from the game. Second,
the considerations just stated will show also that the attributes themselves
are infinitely divisible. Our concept of “elementary attributes” implies no
atomicity. Third, there can exist no absolute measure of similarity (i.e.,
matching correspondence) between non-identical sets of attributes which are
infinite, unbounded, and unconstrained.
Infinity is no simple subject, but we are now trapped into considering
some consequences of invoking it. We find that measures of matching can
in fact exist between certain infinite sets. Let us consider, for instance,
the set of all positive integers A: {1, 2, 3, 4, . ..% and the set of odd
positive integers B: {1, 3, 5, 7, ...} , and regard the elements of these
sets as our selected elementary attributes of two “objects” which being here
equivalent to the sets described, we may also label A and B. Let us then
first take these elements in the order given and, by an act of classification,
Sroupy went ine pare (1). (23), 2 se as “states” of a set G of pluri-state
attributes which we shall call {I, TI, III], IV, ...}. We could define the
‘“nossible” states of the members of G in various ways; one way would
be: the “states” of I comprise the presence of 1 or 2 or 3 o0r...... in the
first position, likewise for II in the second position, and so on. This would
permit the comparison of a jwhole class of sets similar to A and B. Each
of the sets A, B, G is infinite but, in the language of the Cantorian theory
of transfinite numbers, denumerable (Courant and Robbins, 1961; Dantzig,
1962).
If we compare all the attributes, or any ordered sample of them, and
write.1 for a match, 0 for a non-match, we have the correspondence:
Gi sda EI ey TVs ess is
AS 2 Poet Ase rgares
Bis Mee Sie Os (eee rs
giving the matching sequence: 1 0 0 0 ...... , that is, a simple
matching coefficient (Sokal and Sneath, 1963), if we start from 1, of
me matches cael
~ matches--non-matches n
Siip ?
where n is the number of attributes being measured. Obviously, as n tends
to infinity, Sy, tends to zero. If we start elsewhere we have S,, =0 in any
case. This is not very helpful but, with what follows, it Ulustrates the point
20 THE QUEST FOR AN OPTIMAL TAXONOMY
that it is not only the attributes used, but also the way in which they are
grouped and arranged, which affects the results.
If we now look at these sets in another way (one of an infinity of
ways) and take as our set H of two-state attributes {i, ii, ili, iv, ...} the
presence or absence (anywhere in A or B) of the numbers 1, 2, 3, 4, ......
(this is in effect establishing a new set B’ from the union of B with the set
{ absence of 2, absence of 4, ...... }), we have the correspondence:
ys CESS HA Te a Vel 2 DR ea
ADS He i teh ns wae Mee rete
Bacal
Then, by taking the whole or any ordered sample, we have, scoring for
matches as before,1 0 1 0 1 ...... if we start with an odd number, or
Oe Ib D> Ty a coicmdig tic if we start with an even number, giving S,, =4 for
any ordered sample of an even number of attributes from H and S4y +4 (as
the sample size increases) for any ordered sample of an odd number of
attributes from H. We can indeed say that S,, =4 for the comparison (made
in this particular way) of the two infinite sets of elementary attributes.
Moreover, we can say that statistically the most probable estimate of Sy,
from any random sample from H is also 3,* and can in fact evaluate the
probability of obtaining this or any. other result for a sample of any given
size. We have, in fact, for our defined sets and procedure, a parameter S,4,-
and a probability distribution for sampling error.
The infinite sets A, B, and B, just discussed, are in fact (if considered
as linearly ordered sets) bounded in one direction and not in the other, but
it would not affect the argument if we took the sets of all integers and of
all odd integers (positive, zero, and negative), which are not bounded in
either direction. What is more, we may calculate the matches between pluri-
dimensionally ordered sets, such as the two-dimensional arrays:
WRC hele eS eel Dh ORO ES By Mn ich +)
OPI aS ouieyet ga it gD. Ae £3 glues
8) A Be ay 6 ands 444 9505 OialG alt
C4) 3.158 ens Dy 208) 23 bee mn a
vee 6.61 Wa oute |
(a oe Oi ee De O. Reot. RS |
fe eee
giving the “obvious” matching coefficient of Sc, =4, though by considering
other attributes derived from these sets (e.g., presence or absence of a prime
number in a given position) other similarities would be obtained. Such
comparable arrays may be bounded or unbounded in any direction, provided
that they are conformable or can be rendered conformable. If there is no
‘natural” bound, one must of course choose appropriate or arbitrary starting
points for a matching procedure.
Thus, on such a model, having set up a particular correspondence and
using any particular measure of similarity, there does exist an actual para-
metric value of similarity over these infinite sets which we may hope to
estimate in various ways. Why, then, do I say that in the general case of all
attributes, no such parameter exists—in other words, that no meaning can
be attached to the concept of such a parametric value? The reason is that
* Strictly, for odd-numbered samples the two most probable values are those two
of the possible values nearest on either side of 4, and these converge on 3 as the
sample size increases.
= L. A. S. JOHNSON 21
our examples possessed underlying (“built-in”’, if we like) regularities or
patterns which pervaded each of two conformable sets and that, furthermore,
we could find a simple and pervading relationship between these two patterns
themselves, even though the sets were infinite. We had, in fact, a set-up case
with several constraints. In the all-attributes case, no such pervading
regularity can exist, since there are no constraints to prevent us bringing
in any number of other infinite sets of attributes (elementary or pluri-state)
to swamp into infinitesimal proportions any regularity existing among some
of the attributes. As a further complication, describable attributes may be
dependent upon the existence of other attributes, for example blood pigments
cannot be compared as between a bloodless animal and one with blood. This
is one aspect of the “no-comparison” problem familiar in the taximetric
literature, and I have alluded to it in the discussion of ordination.
It is abundantly clear. that, because of such fundamental difficulties, any
recourse to an all-attributes concept or to “true” or overall similarity leads
us into a hopeless morass. We can, of course, though not without problems,
arrive at numerical answers purporting to indicate degrees of similarity (or
“distance’”’) if we restrict ourselves to finite sets of attributes or to samples
from infinite sets of certain restricted types, as we have seen. This involves
subjective decisions, as to:
(i) the set of objects considered usefully comparable;
(ii) the domain of attributes which we consider welovenau to our interest
in the objects;
(iii) the “fineness” with which we analyse the features into elementary
attributes (= states) ;
(iv) the establishment of equivalences or homologies between parts of the
objects under comparison; and the consequent grouping of the
elementary attributes into two- or multi-state sets, thus specifying
what we usually term “the attributes” or “the characters” (“multi-
state” here includes “continuously-varying”) ;
(v) the method and intensity of sampling of the objects;
(vi) the method and intensity of sampling of the acceptable sets of
“relevant” attributes ;
(vii) the quantitative or qualitative measures to be used in expressing the
“states” of each attribute (involving an often arbitrary assignment of
working commensurability )* ;
(viii) the measure of similarity (or distance) to be adopted.
Only some of these are susceptible to a more or less quantitative approach ;
in those cases statistical or other mathematical principles and techniques
will sometimes assist in the judgement but subjective, though not always
arbitrary or uninformed, decisions as to appropriateness or usefulness are
needed all along the line. I shall mention later some useful bases for such
Judgements but, as an initial restriction, we may reasonably confine our
attention to attributes which show a degree of stability or regularity in the
imdividual objects over the time-range in which we are interested. Such a
restriction must itself be arbitrary; furthermore, since it does not in itself
* Williams and Dale (1965) have recognized “the highly autocratic nature’ of the
convention that attributes are dimensionless (‘“dimension” is used here as in dimension
theory of physics, not in a spatial sense) and that quantities representing different
attributes are jointly available for arithmetical manipulation. ‘Standardization’ to
unit variance, or in some other way, may add some reasonableness to the commensur-
ability assumption but does not remove the difficulty; the particular scaling thus derived
depends in any case on the constitution of the chosen sample of objects.
2, THE QUEST FOR AN OPTIMAL TAXONOMY
introduce sufficient constraint to prevent the swamping of overall regularities
by additional attributes, there is still no parametric value of sumilarity.
The concept of a “general” classification has been criticized also by
Edwards and Cavalli-Sforza (1964), while some authors (e.g. Olson, 1964)
have explicitly but unjustifiably assumed that there exists a finite set of
“meaningful” characters.
To resume the enumeration of Gilmour and Walters’ principles:
(7) “General-purpose classifications can be made only when the
objects concerned are influenced by a powerful factor, which causes a
number of their attributes to be highly correlated in their occurrence;
in the absence of such a factor, only special-purpose classifications can
be made.”
Comment: It will be best to avoid discussion of causality; we have
perhaps already ventured too far into the philosophical chamber of horrors!
In discussing the previous principle I stressed the importance of pervading
regularity or pattern (terms which are free of the causative or the particular
mathematical connotations of “factor’’). We need to know not only some-
thing of the patterns but also the range of. their pervasiveness and the extent
of their relevance to our particular concerns. High correlation (or any other
measure of covariation) of occurrence of attributes (i.e., of “states”), which
Gilmour and Walters consider to be the sine qua non for a general classifica-
tion, is meaningful only over a restricted domain, as we have seen, and is
therefore dependent on our interest and purpose. If we attempt a “general”
classification, for example, of the object set:
{Mao Tse-tung, a peanut, the Sphinx, an electron, a litre of alcohol,
the star Achernar} ,
we shall find it very unsatisfying because we cannot define the field of our
interest or readily choose attribute sets showing regularities. Such sets could
be found among the infinity of attributes, but would appear to us to be
absurdly chosen.
The question is whether one can choose such a domain pervaded by a
regularity which, firstly, is perceivable by us, and secondly, seems meaningful
and useful to us. For a set of miscellaneous everyday objects there are
various ways of selecting a set of many pluri-state attributes which would
show a high correlation of their states. For instance, the selection could be
such that a classification derived from a randomly chosen subset of the
original attribute set would have a high probability of grouping the objects
according to their geographical origin, which would then be the “factor” or
regularity determining the high correlation. Our immediate response is, “But
this is a special-purpose classification, and the attributes were specially
selected.” This is true, but when we select (and we do select) the acceptable
attributes to be drawn upon in making a “general” taxonomic classification
we also unconsciously choose a particular set of attributes specially reflecting
a “factor” which is strongly linked to our (selected) interests.
In biological taxonomy, many regularities are discernible among the
attributes which we customarily consider useful for classification, of interest
to other branches of biology, or of practical importance. The reasons for
these qualities and for the regularities are themselves closely though com-
plexly related, and this is the nub of the matter. Evolutionary processes,
selection and adaptation, population structure and dynamics, and the genetic
mechanism itself in its grosser aspects as well as in its physico-chemical basis
and its organization for the storage, replication, transfer, and implementation
of instructive information—all these, intricately interwoven as they are, are
L. A. S. JOHNSON 23
closely associated with those perceivable (phenetic) characteristics of:
organisms which concern us. How this bears on our original questions
concerning optimal classification will be discussed after dealing with a few
remaining points.
“(8) A distinction can be made between typological and definitional
methods of making a classification. In the former, which is exemplified
in the semantic development of the words used in everyday language,
no one or more attributes are necessarily possessed by all the objects
in a particular class, but rather, these objects show a ‘family resemblance’
to an imaginary ‘type-representative’ of the class. The definitional
method, on the other hand, involves a conscious laying-down of certain
attributes that an object must possess in order to belong to a particular
class. Each method is appropriate to a particular type of purpose.”
Comment: Provided that we define the attributes and attribute-sets,
this is a valid methodological distinction. The use of “typological” here may
be misleading, however, since it suggests the essentialistic concept of an
archetype or Bauplan which has been effectively criticized, for example, by
Simpson (1961) and Hull (1965). It will suffice to quote with approval
Simpson’s dicta that “Typological theory is linked with philosophical idealism
[i.e. essentialism. L.J.] which on pragmatic grounds (if no other!) must
be excluded from modern science” and that “such metaphysical beliefs
have no heuristic value’. This is not to deny the usefulness of a sort of
“working” Type concept derived as an ordering of our experience and used
for the purpose of orientation of further experience (Carolin, 1967). Such
a concept. must be subject to any necessary revision in the light of new data
or of reconsideration, and need have neither essentialistic nor phylogenetic
implications. (This has no present connection whatever with the system of
nomenclatural “types” which are now merely formal reference points for the
names of taxa.) The first of the two methods distinguished by Gilmour and
Walters may be redefined to exclude any notion of a “type-representative”
and then becomes in effect the polythetic method (Sokal and Sneath, 1963;
Williams and Dale, 1965) used either intuitively or numerically in most
biological systematics today. The second is similar to the monothetic method
employed by some “old-fashioned” systematists in placing their taxa, and
used generally in keys and some other post-classification procedures for
discrimination and assignment, as well as in ecological, non-biological and
ad hoc classifications. Monotietic classifications are by nature arbitrary or
suitable only for special purposes, and will not henceforth concern us.
Apart from these “Principles”, several other statements of Gilmour and
Walters are pertinent as expressions of the phenetic school’s viewpoint. They
will serve as convenient pegs (not, I hope, as a gallows!) upon which to
hang some observations. Gilmour and Walters refer disapprovingly to ‘the
impression that taxonomic work has an aim of its own, apart from the aims
of biological science as a whole, and that, if this aim could be fully accomp-
lished, it would result in a single, perfect, ideal classification of living things”,
and proceed to uphold “the view that neither affinity nor phylogenetic
relationship can be regarded as valid aims for taxonomy and that they have
been adopted because of a lack of appreciation by biologists of one of the
basic principles of classification enunciated by philosophers—namely the
principle that it should serve some extraneous purpose.”
We may grant that the quest for a single, perfect classification is beset
with difficulties, some of which are inherently insoluble. But what do the
words “should” and “extraneous” mean in this passage? We have seen that
classifications can only be judged in relation to some set of requirements
24 THE QUEST FOR AN OPTIMAL TAXONOMY
(not necessarily an aim of the maker of the classification) but the expression
of phylogeny is as valid as any other requirement, even if it cannot be
perfectly fulfilled. Biological classifications are useful precisely because of
circumstances inextricable from those involved in evolution in general and
the phylogeny of the groups concerned in particular. Hence one cannot speak
of “extraneous” purpose as if this must exclude phylogenetic expression.
‘The authors proceed: “This lack of appreciation has left, so to speak,
a vacuum which was filled, in pre-Darwinian times by the semi-theological
concept of affinity, and later by the equally vague [My emphasis. L.J.] con-
cept of phylogenetic relationship.”
I am at a loss to understand how even a philosopher, much less two
biologists, could make this astonishing statement! Phylogenetic (cladistic)
relationship, though not metrically expressible in any unequivocal way, is
(potentially) topologically representable with little ambiguity, at least above
the level of the coenospecies. We may not know the details of phylogeny
but (unless we reject biological evolution) we must accept that they exist
uniquely in space and time, and therefore form a concrete basis for concepts
of phylogenetic relationship, however defined. In contrast, the notion of
“affinity” is subject to unlimited variation and any claim for a firm basis
for it must be metaphysical. The province of metaphysics appears to be to
pose and seek to answer questions which are either practically or inherently
unanswerable. If the former become practically answerable then that part
of the subject moves into the realm of natural science. The latter have no
place in science—or perhaps in any useful inquiry—except as a warning.
Gilmour and Walters then suggest that the purpose of taxonomy should
be to make a “broad map of the diversity of living things which, by taking
account of as wide a range of attributes as possible, will serve the needs
of aS many as possible of those concerned with animals and plants”; they
regard phylogeny as the “factor” making such a general-purpose classification
possible.
This procedure provides no definable basis for agreement or for testing
whether a classification should be changed or not, except that these authors
would “regard stability of nomenclature as a very important factor in deciding
whether or not to alter the rank of a taxon”. Now, under the present system
formal changes merely of rank (as distinct from transfer from one group to
another) are not called for by changes of position on the topological map of
inferred phylogeny (some palaeontological cases apart) and in fact usually
arise from re-evaluation of the importance of phenetic differences, so that it
is hard to see how the phenetic philosophy itself affects this situation!
From their next remark that “where new knowledge renders an existing
classification clearly absurd, changes must be made, but we suggest that such
cases should be comparatively few”, it is clear that, just like phyleticists,
Gilmour and Walters cannot stomach classifications which are manifestly
inconsistent with the inferred phylogeny or at least with the inferred genetic
constitution. In a later paper, Walters (1965) expresses an even more
extreme pragmatism; one may share his impatience with nomenclatural
formalities and quibbles, but these are not a product of phyleticism.
The virtue of arbitrarily imposed stability of any unsatisfactory system
is not apparent, though frivolous changes should be avoided. Systematics asa
humble servant supplying changeless tags for gardeners, agriculturists,
foresters, or physiologists would contribute little of interest to biology.
Caution, criticism, and common sense play useful roles in science but con-
Servatism for its own sake is inimical to it. If taxonomy were to have only the
uninspiring goals set up for it by Gilmour and Walters it might as well be
L. A. S. JOHNSON 25
abandoned to any rude mechanicals who cared to gather voluminous data for
a computer. Understanding of nature would not be its purpose and any value
it might have for biological theory would be incidental.
Tue Victous CIRCLE
Another philosophical defence of phenetics rests on the assertion that
to allow phylogenetic conclusions (speculations, hypotheses) to influence
taxonomic conclusions involves us in circular reasoning and is therefore
inadmissible. This view is forcefully expressed by Sokal and Sneath (1963)
and accepted by Williams and Dale (1965). Many practising systematists
(e.g., Carolin, 1967) now defend it in theory, but nevertheless appear to be
influenced by phylogenetic thinking in their actual work.
It is true that there are some recursive loops in the phyletic taxonomist’s
reasoning. This applies equally to morphological interpretation and descrip-
tion, in particular to the problem of homology. Attempts to develop “logical”,
non-phylogenetic definitions of homology (e.g., Mason, 1957; Carolin, 1967)
are doomed to failure, being open to the same criticisms as phenetic
taxonomy.* Indeed homology needs no separate treatment since comparative
morphology implies taxonomy and vice versa. The need for very careful inter-
pretation of morphological homology on a basis of evolutionary likelihood is
illustrated by the highly modified and cryptic structures of many animals
and plants, for instance the “bulrush” or “cat-tail” genus Typha (Briggs and
Johnson, 1968) as considered in relation to other groups of monocotyledons.
No phenetic comparison could be of much value here unless the relevant
homologies were for the most part correctly worked out beforehand. The
comparative survey by Hamann (1961) of the families of the “Farinosae’”,
though most comprehensive and thoughtful, includes a numerical phenetic
study of which we cannot confidently accept the results, because of serious
doubt as to the homology of many of the features compared. Here, as often
in broad surveys, the data have been drawn partly from various published
descriptions, which may be very misleading. Throckmorton (1965) points
out that phenetic parallelism may be equivalent to genotypic homology, that
is, Similar conditions which appear to arise independently may be due to
separate “assembling” of elements of a common pre-existing genotype.
We have seen that there is a hidden circularity in any definition of a
“sveneral classification” and a hidden infinite regress in every aspect of
phenetics. The Infinite Regress and the Vicious Circle or ultimate tautology
are the two most inescapable hard facts of philosophy, mathematics, and
science—at bottom we must come upon one or the other. They have been
encountered by every critical child who has asked, “If God [or whatever
agent or process one wishes to substitute] made the Universe, then who made
God?” In any case, what is before or after, backwards or forwards, on a
cosmic (or ultra-cosmic) scale? Such ancient questions are ultimately
* Key (1967) attempts to define operational homology thus: ‘Feature a, of organism
A is said to be homologous with feature b, of organism B if comparison of a, and b,
with each other, rather than with any third feature, is a necessary condition for
minimising the overall difference between A and B”. Despite its spuriously precise
formulation, such a definition is itself thoroughly non-operational except in finite cases,
for the reasons already given. When discussing set-correspondences, I have. used the
term “working homologies” simply for such correspondences of features (of objects
of any kind) as we may take, possibly on very complex and subtle grounds, to be a
reasonable basis for further steps in the comparative procedure. The “operational
homology” concept discussed at length by Sokal and Sneath (1963) boils down to much
the same thing; it is only vaguely operational, in the sense of having some empirical
consequences which can be checked (Hull, 1967).
26 THE QUEST FOR AN OPTIMAL TAXONOMY
unanswerable, though they may be pushed back somewhat along the regress,
but they may help us to dispense with some unnecessary non-answers to non-
questions.
The partial circularity inherent in the phylogenetic approach is not
reducible to a petitio principi as is the simple vicious circle of logic. It is
not just saying tautologically that conclusion B follows from premise A when
we have already used B in formulating A (as, for instance, “a classification
produced by a good taxonomist is good”, having defined a good taxonomist as
“one who always produces good classifications”). Nor is it arguing in an
epistemological circle (Hull, 1967), that is, purporting to show that B is
true because of A when we can only know that A is true if we know that B
is true (as, for instance, “a newly discovered mammal from Africa is a
placental mammal because all African mammals are placentals”).
Rather, we are accepting a degree of positive feedback in what is not a
chain but an anastomosing plexus of reasoning and evidence. The premises
and reasoning used in arriving at a representation of a (hopefully) deduced
phylogeny are highly complex, as are the conclusions (see, for example, the
attempted reconstructions of the phylogeny of Dipsacaceae by Ehrendorfer,
1964a, 1964b, and of Proteaceae by Johnson and Briggs, 1963; also Thorne,
1963). There is no hope of making them fully explicit, but to deny them
validity on that ground would be to deny validity to all but the simplest
reasoned conclusions in science or life. Many pheneticists take the view that,
in the absence of a “time-machine” (Carolin, 1967), palaeontological data
provide the only information of direct phylogenetic significance that can be
validly fed into a taxonomic system. But most of us do, after all, believe
that the processes of evolution are pretty well understood. Most phylo-
genetically inclined taxonomists and general evolutionists consider that the
existing corpus of evidence justifies the use of evolutionary principles to make
sense of the diversity of Recent as well as fossil organisms. This involves
a great deal of argument by analogy and extrapolation, but so does most of
science. How many of us know from direct evidence that we consist of
neutrons, protons, and electrons, or even that we have chromosomes? But
we believe it because we have convincing evidence of consistency in the
physical world. In the course of describing and analysing the phylogenist’s,
and in fact the common scientific, method of successive approximation, Hull
(1965, 1967) presents powerful philosophical arguments for a similar rejec-
tion of the charge that phyletics is based on viciously circular reasoning, in
either the logical or epistemological sense.
It is a commonplace that, outside mathematics, most scientific hypotheses
and theories are inductively derived, though we aim to reduce the number of
such hypotheses to a minimum and to use them as postulates for a deductive
superstructure. They are testable by predictive value, self-consistency, and
consistency with extraneous evidence. Operationism requires a hypothesis to
be potentially falsifiable, that is, subject to tests which could show it to be
false. If we exclude phylogenetic interpretation, phenetic “general’’ classi-
fications of organisms are scarcely hypotheses at all*; they are subject merely
to practical evaluation by their predictivity within the field of our interest.
Phyletic classifications (which of course use phenetic techniques as part
of their method of derivation) may certainly be equally effectively judged
* This is admitted by Williams and Dale (1965), though we may accept their
suggestion that non-probabilistic numerical classifications in other fields have value as
hypothesis-generating systems. While they allow that phylogenetic hypotheses may be
generated, these authors tend to dismiss them on the ground of their alleged
untestability.
— O°
L. A. S. JOHNSON Pat
on predictivity; they also embody biological hypotheses (though the classi-
fication as such may not fully express them), which are subject to many
checks of self-consistency and. extrinsic consistency—though such checks are
not necessarily easily expressed in quantitative terms. It has been claimed
(e.g., by Birch and Ehrlich, 1967a@) that these embodied hypotheses, and
indeed synthetic evolutionary theory generally, are not falsifiable. This is
due to the unsophisticated assumption that a useful hypothesis must be a
sufficiently simple and definable proposition to be falsifiable in the same way
as the statements: “all swans are white” (known to be false) and “the
equation #£"+y"=2" has no solutions where 2, y, 2, and n are integers and
n > 2” (the celebrated “Last Theorem” of Fermat, which is not a deductively
proved theorem at all but has not been shown to be false). Phylogenetic
‘hypotheses are susceptible of demonstration that they are very likely to be
untrue, and this is indeed the only disprovability we can hope for in a good
deal of science. We can achieve a considerable and reasonably convincing
understanding of nature in such ways, and can apply a great deal of criticism
to our evidence, arguments, and conclusions. Mackerras (1964), in a cogent
defence of the phylogenetic method, says that “to make a preliminary arrange-
ment on general resemblance (as is often done), test its components for
phylogenetic concordance or discordance, and then base a classification on
the results of those tests, is not circular reasoning in any sense of the words.”
Birch and Ehrlich (19670) say, “We do not need to consider the unknown
history of the organisms .. . in order to classify them . . . Phylogenetic
history ...is not pertinent to most of the uses of classification. Astronomers
do not have to know the histories of sidereal bodies, nor physicists of atomic
particles in order to do valid scientific work with them.” These authors claim
precision for phenetic methodology and are opposed to “mixtures” of phen-
etics and phyletics in classificatory procedures. An outrageous stab on their
part is to accuse the evolutionist Ernst Mayr of Platonism; to interpret
Mayr’s insistence on the “reality” (i.e., reasonable definability) and import-
ance of the species in evolutionary theory as a form of Platonic essentialism
is a gross distortion indeed. Birch and Ehrlich’s quoted statement is partly
true; we certainly can classify without reference to phylogeny (Linnaeus
did it!) and, for the reasons already given, such classifications may work
quite well. This is in itself no reason at all to reject consideration of
phylogeny, which is one important aspect of the evolutionary process that
underlies the workability of our classifications. Astronomers do, of course,
consider the histories of stars and may modify their classifications accordingly.
The case of [sub]atomic particles is simpler and not really comparable;
nevertheless elucidation of underlying regularities and the possible trans-
formations of particles is highly pertinent to our mental systematization of
them. Improved classification in these cases does in fact involve positive
feedback (i.e., a “mixed” procedure).
For the reconstruction of phylogenies we can use evidence of many kinds
which is not just derived from the patterns of phenetic diversity we observe
in the organisms under study. Cytogenetic data, for example, have a special
relevance beyond mere matching of attributes of the chromosomes or of the
genetic systems. The contribution of comparative biochemistry, especially
where its adaptational significance is evident or where biosynthetic pathways
indicate probable directions of change (e.g., Scora, 1967), frequently goes
beyond mere phenetic comparison. Quite often there is a fossil record of
considerable value. In particular, probable directions in the evolution of
organ-systems can be deduced from ontogenetic, ecological, adaptational, and
historical evidence and comparisons. Examples are legion and can be found
28 THE QUEST FOR AN OPTIMAL TAXONOMY
in works on evolutionary theory and in many publications presenting critically
argued reconstructions, as in the case of the Proteaceae, already cited. A
single example will suffice: floral adaptations for pollination by long-tongued
insects or by birds cannot have occurred in geological periods before such
animals existed. Any phenetic classification which grouped organisms in a
manner inconsistent with such a fact (taking all other relevant information
into account) would not be acceptable as even roughly consistent with
phylogeny, and there is no reason why we should be asked to accept it
simply because of claims of repeatability, objectivity, precision, or stability.
Repeatability is not dependent on the phenetic approach and the claimed
objectivity and precision are superficial. Stability has been mentioned already
and later will be discussed further.
In summary, such philosophical considerations as we have reviewed, far
from invalidating the phyletic approach to taxonomy, may be turned against
the pheneticists themselves.
TAXIMETRICS*. J. NUMERICAL PHENETICS
Evaluation and methods
We have discussed at length the foundations of phenetics. Only brief
mention can be made of the techniques of numerical taxonomy and their
applicability. These are being used, often on rather trivial problems, by
increasing numbers of taxonomists who often seem to accept very uncritically
the philosophical foundations and the mathematical models which they
explicitly or implicitly embody. The worth of the results seems usually to
be measured against what the taxonomist has done, or would have done,
without the techniques—an amusing test, to say the least. Mathematical
methods of ordination and classification seem to have thrown very little
new light on the taxonomic problems of “difficult” groups, despite their use-
fulness in ecology or in non-biological fields where there is no obviously
prime source of regularity comparable with phylogeny and the genetic
constitution of organisms.
It is true that Watson et al. (1966, 1967) have produced a slightly better
classification of the Epacridaceae and a considerably better one of the
EKricaceae than those previously existing (not necessarily “accepted” as they
put it)—but the latter were almost a century old and based on less complete
data and, in the case of Drude’s less satisfactory classification, on an analytic
or monothetic approach. From the data, it seems to me that many good
modern taxonomists should have been able to effect equal improvement by
adequate study and phylogenetic consideration. Indeed, by careful study
and reasoning, Watson had already cleared up most of the problems, though
the computer analysis suggested some placings for doubtful genera. In the
more difficult case of the Basidiomycetes, reported by Kendrick and Weresub
(1966), “Adansonian” computer analyses gave thoroughly unsatisfactory
results as compared with a reasoned phylogenetic approach. It would be
difficult to explain these away by criticism of the mathematical models or
*The forms “taxometrics” and “taxonometrics”’ are also in current use. “Taxi-
metrics” is etymologically the best-formed since the first root is the Greek rdéts, ragews
(or Ionic rdééos)—‘‘an arranging”, of which the combining form in Greek compounds
was raé-(Liddell and Scott, 1864). The word ‘taxonomy’ is badly formed (cf. the
French “taxinomie”’) but is now firmly established by usage. One may hope that _
“taximetrics”, as used by Rogers (1963), will prevail, or perhaps it is not yet too late
to substitute the simpler “taximetry’, with the ending on the model of “geometry”,
“trigonometry”, “biometry”, “anthropometry”, etc. Further development of the first of
these examples could lead to the delightful consequence of numerical taxonomists’ being.
called ‘‘taximeters”’.
L. A. §. JOHNSON 29
techniques employed; the source of trouble seems to have been evolutionary
convergence. In a group of Solanwm species and hybrids, Heiser et al. (1965)
found that the results of a phenetic numerical analysis were less satisfactory
than those of a “subjective” study when checked with the considerable bio-
systematic information.
A recent coup @ewil. over the grass family, using a pr opaiilisere mixed-
data method (Clifford and Goodall, 1967), seems only to confirm the well-
established features of modern non-numerical classifications and to indicate
that the “difficult” cases actually are so. Its sampling basis of one species
per tribe would certainly satisfy neither traditional taxonomists nor
phyleticists. The authors claim it as an “advantage” that “such a limited
sampling eliminates all the intra-taxon variability”, which hardly engenders
confidence in their appreciation of the significance of nature’s complexity.
In their study of difficult species-complexes in Cassia, Irwin and Rogers
(1967), using the graph-theory model of Wirth et al. (1966), seem to be
reasonably well satisfied with the results. The taxa are too closely related
to show clear phylogenetic patterns, but little biosystematic information is
available. The taximetric clustering procedure seems at least to have
promoted the authors’ confidence.
A notable exponent of numerical methods recently remarked to me that,
given suitable data and an appropriate program, “the computer can produce
a better classification than a poor taxonomist”. But then, so can a good
taxonomist—and, being in possession of much extraneous information and
reasoning power which are not in the storage unit or the programmed
strategy of the computer, he can often do so with much less tedious recording
of data. Might it not be more economical to employ a few more good taxo-
nomists? This ought to be possible out of a world population of 3 x 10°.
The literature provides ample illustrations of considerably divergent
classifications of the same material, from the same data, produced by various
strategies of numerical taxonomy (Boyce, 1964; Olson, 1964; Katz and Torres,
1965; Minkoff, 1965; Rohlf and Sokal, 1965: Sheals, 1965; Kendrick and
Weresub, 1966; Lance and Williams, 1966a@; Sokal and Michener, 1967;
*t Mannetje, 1967). Varying the attribute sampling gives somewhat different
results again; so does changed scoring, whether it implies altered attribute
definitions or grouping, changes in measure, or all of these. It is true that
different methods sometimes yield reasonably concordant results for the major
divisions of a group but in these circumstances the taxonomist is seldom in
doubt in any case. If, after a careful investigation, a taxonomist remains in
doubt as to the classification of particular groups, it is usually due to intrinsic
complexities, past or present, in evolutionary situations. Lack of well-defined
clustering and especially of nested groupings is inherent in such situations;
consequently the different characteristics of the various taximetric strategies
will result in lack of consistency. These instabilities arise from the funda-
mental difficulties discussed earlier; they cannot be regarded simply as
functions of a signal : noise ratio (Lange et al., 1965) (though this may vary
with the method), since the signal itself is to some extent what we define it
to be.
The theoreticians of numerical taxonomy have enjoyed themselves
immensely over the past decade (though not without developing several
schools with scant respect for each other!). The mushrooming literature is
quite fascinating and new developments tumble after each other. Anyone who
is prepared to learn quite a deal of matrix algebra, some classical mathe-
matical statistics, some advanced geometry, a little set theory, perhaps a
little information theory and graph theory, and some computer technique,
30 THE QUEST FOR AN OPTIMAL TAXONOMY
and who has access to a good computer and enjoys mathematics (as he must
if he gets this far!) will probably find the development of new taximetric
methods much more rewarding, more up-to-date, more “general”, and hence
more prestigious than merely classifying plants or animals or working out
their phylogenies. Unlike the taxonomic questions themselves, the methodo-
logical questions which the taximetric mathematician sets himself are mostly
so definite and so angswerable—yet hunting for the answers is difficult enough
to be interesting:
Does a certain distance function define a metric, semi-metric, quasi-
metric, or non-metric space? Is it monotonic (varying throughout in the
same direction) as object-groups are successively fused? What are the
properties of the Canberra metric as a variant of the Manhattan metric?
What are the relative advantages of divisive and agglomerative strategies?
(One can do a little classifying of the strategies themselves—without using
the computer, so that there is some intellectual satisfaction in it.) What are
the properties and advantages of hierarchical (nested) systems as opposed
to overlapping, clustering (clumping) systems? To what extent are particular
agglomerative sorting strategies space-distorting? Can dimension-reducing
ordination techniques such as principal component analysis and factor
analysis be used efficiently as a basis for extracting a classification from an
ordination? What are the virtues of rotation of axes in a factor analysis to
yield “simple structure’? What do the “factors” of factor analysis mean?
(Not so easy to answer.) Are certain matrices always positive semi-definite
(symmetric with all eigenvalues non-negative)? Are certain functions
algebraically tractable? Do annoying singularities occur? Are probabilistic
models really applicable or desirable? Can predictivity be usefully measured ?
What are the virtues of combining R- and Q-analysis, and what are the
appropriate techniques? (As now agreed both by Sokal and Sneath (1963)
and Williams and Dale (1965), these terms refer to matrices: the elements
of a Q-matrix are measures of comparison of objects (individuals) while those
of an R-matrix are measures of association of attributes.) How effective are
heuristic “hill-climbing” strategies (Rubin, 1967) which search for optimal
structure by systematic trial? How large a data matrix can particular
computers handle for particular programs and how much computer time
does an analysis take? Are programs easily modifiable? What methods can
handle mixed data, and how validly? Should attribute scores be standardized
to unit variance, or in some other way? Are double-negative matches to be
included? What is negative in any case? (Suppression of a character may
well be the derived condition within our frame of reference; to assign greater
importance to “presence” than “absence” is a decidedly subjective judgement. )
Is the information-statistic the best basis for agglomerative strategies?
Some of these questions are discussed in an enlightening paper by
Williams and Dale (1965). Although these authors make the usual, partially
invalid, pheneticist assumptions, their mathematical discussion is most help-
ful and pertinent. However, reference to various subsequent papers is neces-
sary (e.g., Bonner, 1964, 1965; Macnaughton-Smith, 1965; Hall, 1965, 1967 a,
19676; Davidson and Dunn, 1966; Goodall, 1966; Gower, 1966; Jancey, 1966;
Menitskil, 1966; Wirth et al., 1966; Lance and Williams, 1966a, 19666, 1967a,
1967b, 1967¢; Davidson, 1967; Estabrook, 1967; Orloci, 1967; Rubin, 1967;
Crovello, 1968; Wallace and Boulton, 1968), and indeed any attempt to be
comprehensive in this field is obsolescent before it reaches the printer. The
well-known text of Sokal and Sneath (1963), which was welcomed by non-
initiates as the Bible of the subject, though still important, has alre ady taken
L. A. S. JOHNSON 31
on the archaic flavour of an Old Testament, both as to foundations and as to
the range and evaluation of mathematical methods and models.
Are phenetic numerical methods, then, of value in practical systematics?
I think they can be, especially now that computers can process high-order
matrices (approaching 200 “objects”, or even more) and that mixed-data
programs have been developed to deal with two-state, multi-state, and infinite-
state (continuously-varying) quantities in the same matrix (Lance and
Williams, 1967¢; Wallace and Boulton, 1968). Our ordinary intuitive pro-
cesses begin to lose efficiency with problems of this magnitude, especially
when a comparable number of attributes is used. The computer’s elucidation
of “structure” in the data may be useful to us in suggesting an appropriate
classification, even though we are aware of the numerous subjective and
arbitrary decisions implied in the choice of data and of mathematical models.
I do not believe, however, that we should accept any such classification
as the last word or as indicating that answers exist to the chain of questions
posed early in this address. Ordinarily one can attempt to evaluate such a
classification by evolutionary considerations and can modify it accordingly
if necessary—there is no need to stand in awe of its “objectivity”. If this
is not feasible, one can accept it as possibly better than one could do by
intuitive methods. Having accepted a finite set of relevant attributes (not
necessarily all wsed in the analysis), one may judge classifications as to
predictivity within that set by means of the probabilistic utility function of
Goodall, who, in an interesting paper (Goodall, 1966), outlines some of the
constraints which are necessary before any probabilistic technique can be
validly used.
Warburton (1967) suggests that the purpose of classification should be
to maximize “the probability that statements known to be true of two
organisms are true of all members of the smallest taxon to which they both
belong. It should not be impossible to develop tests for this property to
objectively decide which of several rival classifications is best.” Any such
test, of course, would depend on definition of a finite set of “statements”. If
we agree to that, Warburton’s criterion could provide a useful pragmatic
test within a frame of reference, but because of this arbitrary aspect it
answers no general question about the best classification.
No good reason exists why any particular horizontal cuts across pheno-
grams* should be accepted as meaningful “phenon levels” (Sneath and Sokal
1962; Sokal and Rohlf, 1962; Sokal and Sneath, 1963) upon which to erect our
formal taxonomy. All claims for phenetic “standards” of rank collapse on
analysis, including those reiterated in the rather brash paper in which Sokal
and Sneath (1966) set out their recipes for a great leap forward to “efficiency
in taxonomy”.
Although a number of earlier authors had proposed numerical methods
for taxonomic classification, it is interesting that one of the influential
streams in the modern period is associated with the work of P. H. A. Sneath
(for references see Sokal and Sneath, 1963) on the classification of bacteria,
a group in which phylogenetic interpretation has been unsatisfactory and
equivocal. Another vigorous stream, associated with W. T. Williams and
his collaborators, began in the field of ecology and has close associations
with workers in such areas as criminology (Macnaughton-Smith, 1965),
industry, and business, in all of which classification has been and must
* Phenograms are phenetic dendrograms purporting to show similarity by a tree-
like diagram of nested subsets of the objects—not merely topologically but scaled
according to the particular measure used.
By THE QUEST FOR AN OPTIMAL TAXONOMY
surely remain an ad hoc or arbitrary matter (however “objective” we may
cause it to appear by making the subjective decisions before beginning the
calculations ).
Generally experienced systematists, young or old, who employ taximetric
methods usually seem to make excuses for them: “Well, the computer analysis
didn’t do any better than (or as well as) I could, but it wasn’t a bad job and
perhaps it gave me an odd idea here and there” would sum these up. Despite
substantial response arising from genuine interest as well as from band-
waggoning, the showing after ten years’ hard selling is not at all impressive
so far as improved practical classification or biological understanding are
concerned. The onus must surely be increasingly on the numerical phenetic-
ists to give reasons, other than its meretricious glitter, why we should buy
their product except with considerable reserve. Fashion, spurious objectivity,
and competition for financial grants (Rollins, 1965) are not very scientific
reasons. Neither is fear of being included by such determinedly iconoclastic
zealots as Paul Ehrlich (1965); in the class: { “members of the old school”
who “would still like to see a pinch of phylogenetic speculation mixed into
their basic data (presumably for sentimental reasons)” and who “will con-
tinue to promote this confusion for some time to come”} —and thus excluded
from the class: { “those who wish to look forward”} , regarded by Ehrlich
(objectively, no doubt) as non-overlapping with the former class. There is
a familiar ring to this: “He who disagrees with me is a reactionary.”
Our only certain scientific pay-off from phenetic taximetric methods is
that, having gathered a lot of data, we shall therefore be less likely to over-
look features of evolutionary or practical significance. Often the game may
not be worth the candle.
Relation to genetic basis
Sneath and Sokal (1962) introduced the concept of the “matches
asymptote”, hypothesizing (Sokal and Sneath, 1963) that “the similarity
between two operational taxonomic units is some parametric proportion of
character matches which we are estimating with a sample of characters” and
that ‘fas the number of characters sampled increases, the yalue of the simi-
larity coefficient becomes more stable’. The meaningless of a parametric
value in the all-attributes case has been demonstrated but it may still seem
reasonable to consider that, given a suitable measure of similarity, such a
parameter exists for the “matches in the nucleotide sequence of the DNA of
the genotype”. If that is so, then, over the attributes ordinarily regarded as
important, the confidence band for an estimate of this parameter will narrow
as the sample size increases. In that there is a certain regularity, related to
the genetic information, over the attributes we are likely to consider, some
such convergence is indeed to be expected. The existence of a numerically
definite asymptote will of course depend on the acceptance of a particular
finite attribute set defined by enumeration of its elements, but it is doubtless
correct to say, in an imprecise way, that the larger the sample of attributes
the better our comparisons will tend to be in reflecting the similarities of
the genotypes. However, even if we knew the entire nucleotide sequences over
a Set of organisms wwe should still have to make many decisions on matching
procedure. (We can hardly speak of comparing “genes”—the gene is no
longer a useful operational unit at the level of molecular genetics.) We
certainly could not set up pluri-unique correspondences of DNA _ base
:
7 More recently (Ehrlich and Ehrlich, 1967), this author has realised that
numerical phenetics is inherently indeterminate in its results——and appears to have
adopted a nihilistic attitude to taxonomic improvement in general.
~ L. A. S. JOHNSON 33
sequences over the whole genotype of organisms which were at all diverse or
differed in karyotype (see Ehrlich, 1964, and Reynolds, 1965, for possible
further difficulties). Taking the step up to the comparison of proteins will
not remove this difficulty, though such comparisons as can be made in this
field will provide information at least as useful as that from other attributes.
Related to, but not identical with, the “matches-asymptote” hypothesis
is the “non-specificity hypothesis” (Sneath and Sokal, 1962; Sokal and Sneath,
1963) which assumes that “there are no distinct large classes of genes affecting
exclusively one class of characters such as morphological, physiological or
ethological, or affecting special regions of the organism”. This suffers from
the same fundamental difficulties of definition and testing as other phenetic
concepts but, so far as the phenetic tests may be accepted as valid, it appears
-often not to hold very well (e.g., Rohlf, 1965; Thornton and Wong, 1967).
Indeed, the lack of correspondence of groupings derived from different sets
of data (say, from internal anatomy and external morphology, or from larval
and adult stages) is a problem in phyletic as well as in phenetic taxonomy,
but at least the phyletic approach to its resolution should be more intelligent
and subtle than crude lumping of the data or results. An alternative, of
course, is to take the view of Ehrlich (1964, 1965) that “phylogenetic specula-
tion is fun, but seems to have little scientific purpose” and hence to regard
all classifications as special, so that “one wishing to make predictions about
the distribution and ecology of larval mosquitoes would presumably do better
to work with a taxonomy based only on characters of the larvae”.
Phylogeny cannot be perfectly elucidated, and it is inherently complex
and reticulate at those levels of grouping at which interbreeding among
groups still occurs. Nevertheless its unique existence provides the nearest
thing to a solid base for general taxonomy.
Taximetrics. Il]. NumMeErRicAL PHYLETICS
- Conditioned as they have become by reiterated assertions to believe that
operationist philosophy demands that taxonomy must be purely phenetic,
only a few of the theoreticians of taximetrics have shown interest in develop-
ing mathematical models and corresponding numerical techniques for the
elucidation of phylogeny. One can set up plausible, though clearly over-
simplified, models of phylogenetic processes, just as one can for genetic
systems. Some at least of these models are reasonably tractable mathemati-
cally, as are the rather idealized models of population genetics which have
often proved fruitful in suggesting hypotheses and in testing for consistency.
The “advancement index” of Sporne (1948, 1954, 1956, 1960) was an early
attempt to assess evolutionary advancement. Sporne’s approach was different
from later methods discussed here, but is of limited applicability, for the
reasons given by Davis and Heywood (1963, p. 39). The concept of a statis-
tically measured overall “advancement” is both too indefinitely based to have
much statistical validity and too generalized for effective reconstruction of
phylogenies.
A simple phylogenetic consistency test, based on the postulate of uni-
directional change in individual characters, was developed by Wilson (1965).
This field of study has been termed cladistics by Camin and Sokal (1965)
who, having formulated a set of assumptions regarding evolutionary sequences,
developed a computer strategy based on the principle of parsimony. Its aim
was to construct cladograms (phyletic dendrograms) representing an evolu-
tionary minimum-path branching pattern for the set of OTU’s under con-
sideration. The method was checked against the palaeontologically well-
(0)
34 THE QUEST FOR AN OPTIMAL TAXONOMY
documented phylogeny of horses and also for a group of imaginary organisms
(“Caminalcules”’) which had been independently generated using the phylo-
genetic principles enunciated. This important pioneering work has naturally
appealed to those phyleticists who do not wish to turn their backs altogether
on numerical methods, but has been virtually ignored by many pheneticists.
Camin and Sokal themselves continue to prefer a phenetic basis for classifica-
tion “until an operational system combining cladistics and phenetics can be
established”; since any such system must be arbitrary, from the nature of the
phenetic component, I cannot see how this aim can be achieved with the
“objectivity” which Camin and Sokal desire.
Cavalli-Sforza and Edwards (1967) have discussed, in considerably
greater depth and detail, some mathematical models and estimation pro-
cedures for phylogenetic analysis. Their intention, only partially fulfilled,
was to use maximum-likelihood methods to estimate the form and proportions
of “the most probable tree uniting the presently living populations”. Their
“branching random walk” model assumes that evolutionary changes leading
to divergences in genetic constitution are, at the level concerned, essentially
stochastic, summarized as “random genetic drift and variable selection”.
One could disagree with these assumptions, and it is clear from the authors’
very honest discussion that many arbitrary decisions have to be made as to
the nature of the spaces and metrics.implied by the models and by the methods
of estimation. Tests of the method in cases where there is good fossil and
other evidence for a particular phylogeny should be interesting. Cavalli-Sforza
and Edwards dismiss Camin and Sokal’s paper with the observation that the
latter authors’ assumption “that evolution proceeds according to some mini-
mum principle’ cannot justify the use of a “method of minimum evolution”,
though Cavalli-Sforza and Edwards do not deny the possible usefulness of
such methods. .
Working with amino acid sequences of cytochrome ¢ from an assortment
of vertebrates, two insects, and three fungi, Fitch and Margoliash (1967) have
used a numerical method of phylogenetic analysis on the basis of “mutation
distance’, that is, the minimal number of nucleotides which must be altered
in order to convert the coding for one cytochrome to that for another. This
is an excellent paper, but I cannot resist quoting the metaphorical gem that
if “one wishes to test a tree which differs only in the order in which the
chicken, duck, and penguin are joined, the only legs in need of recalculation
are those five descending to these birds from the avian apex”. The mental
picture which this conjures up is slightly less surrealistic if we note that
Fitch and Margoliash’s trees grow upside-down and have legs instead of
branches!
Silvestri and Hill (1964) stress the value of the patristic approach (i.e.,
genetic comparison) in microbiological systematics, since there are few
reliable data for cladistic hypotheses in that field. However, Silvestri
(1964) also points out that phenetic differentiation may be outstripped by
DNA evolution because of the degeneracy of the genetic code.
Further development of numerical phyletics is certain and seems to have
considerable promise, but inasmuch as the methods are numerical they impose
their inbuilt metric properties on a situation for which topological “reality”
is more or less inherent, but “distance”, if defined on essentially phenetic
grounds, is dependent on our point of view. “Mutation distance” will usually
be determinable only in respect of a few loci, and therefore inadequately
sampled, but it does perhaps represent the nearest thing to a “natural” metric
basis for biological taxonomy.
we)
Or
L. A. S. JOHNSON
QUALITY AND QUANTITY IN MATHEMATICS
The urge to quantify is upon all biology and the social sciences.
Undoubtedly it has already brought considerable benefits, but we should not
forget that science is not the process of measuring natural phenomena; that
is merely a technique of science. Very largely the aim of science is to discern
qualitative differences, that is, to simplify and reduce the number of quantities
which it is necessary to specify. For instance, in particle physics today the
algebraic theory of groups (this term has here no connection with “groups”,
i.e., classes or sets, in classification) has been successfully applied to the
ordering and prediction of quantum phenomena, as indeed it was applied
earlier to symmetry relations in such fields as crystallography. The essence
of the algebraic concept of a group is not quantity but structure: a particular
set of relations between elements, the latter usually being operations in the
particular class of relations which satisfy the group concept. This principle
is expressed with greater generality in the concept of isomorphism: the
matching of sets of relations, not quantities. It is these qualitative aspects
with which much of modern algebra is concerned (e.g., Maxwell, 1965;
Hollingsworth, 1967). The same is true of much advanced geometry, especi-
ally the non-metric geometry of position (mostly abstract and expressed in
a notation allied to that of set theory): topology, or analysis situs as it was
once called.
We should not be bewitched by number, in particular by the continuum
(for basic concepts and some enlightening philosophical discussion, see
Dantzig, 1962). We shall probably never be able to express or apprehend
complex situations in their precise quantitative detail. Over the centuries,
beginning with the natural numbers1, 2, 3, ...... , we have had to extend
our concept of quantity to embrace zero, directed (positive and negative)
numbers, rational fractions, algebraic and transcendental irrationals, complex
numbers, and: various kinds of hypercomplex numbers, such as quaternions,
vectors, and matrices. To cope with change and with infinite processes, the
differential and integral calculus was necessary. Further, to deal precisely
with matters as uncomplicated (as compared, say, with those of biology) as
the specification of siniple events and relations in space-time, mathematicians
have combined vectors, matrices, and calculus into the formidable subject of
tensor analysis— and this in cases where there are no difficulties of multiple
incommensurability. We have no hope of extending this kind of precise
quantitative mathematics to describe the biological situations encountered
in taxonomy. Mathematics is hard, paradoxically, because its subject matter
is simple—and the subject has thus been able to progress a long way.
Statistics, largely based on probability theory, has been developed to
cope with circumstances, in physics as well as in less simple fields, where
we are concerned with large numbers of events which display a degree of
disorder, that is, lack of individual specifiability (except where individual
observation is feasible). Statistics requires certain assumptions, as we have
seen, which are often not justified in systems of incommensurables. May we
not be well advised to avoid putting all our money on Quantity? Qualitative
assessment (which, as we have seen, does not exclude mathematical concepts)
may be a surer winner in heavy going. To be sure, we can often extract
qualitative structure, or qualitative generalizations, from quantitative data
and, when we can, we should doubtless do so.
CLADISTIC SPACES
We can also very often perceive qualitative structure, and subject our
interpretation to empirical tests, by qualitative means. Quantitative expres-
Sion of a situation, or a quantitative check, is of no greater value unless the
36 THE QUEST FOR AN OPTIMAL TAXONOMY
assumptions underlying it are justified. We now know that often they are
not. In the words of Hull (1965), an author whom all the devotees of
numerical objectivity (and all phyleticists!) should read, “talk of taxonomic
space, like talk of gene pools, is strictly metaphorical. .. . Taxonomic
space ... is an amorphous continuum with no intrinsic metric; there is
nothing about taxonomic space to indicate how long to make the unit of
measurement or at what point to begin measuring once the unit is chosen”.
To this we may add that the dimensionality of taxonomic space, even in the
sense of phyletic space, is arbitrary and the space is not isotropic (similar in
properties in all directions).* There is a time dimension which is metrizable
for our purposes by the ordinary physical criteria. The remaining dimensions
will depend on our model or mental construct of the space. Two main
phyletic cases arise: (I) We can say that there are n dimensions (including
time) where 7 is the number of recognized “ends” (individuals, species, OTU’s
or whatever our smallest unit of taxonomic division is taken to be). This
will be a pure “cladon-space”, and any cladistic tree within it can be mapped
without loss of information on to a two-dimensional space to form a conven-
tional dendrogram, provided that we are not concerned with any measure of
distance or indeed with any ordering except on the time co-ordinate. In this
case it is only the order of branching which counts. (II) We can admit
p+1 dimensions (including time) where p is the number of attributes taken
into account. This class of phyletic-phenetic space, which is merely an
attribute-space with a time dimension added, is the basis of Cavalli-Sforza
and Edwards’s primary model; it has some metric properties but need not
be fully metric in the sense of Williams and Dale (1965). A tree in it cannot
be mapped on to a two-dimensional space without considerable loss of
information. Two- or three-dimensional representations such as_ those
advocated by Sporne (1956), among many others, though useful as illustra-
tions, cannot hope to be very accurate in representing either phenetic or
evolutionary distance, according to whatever measure we adopt.
Whether or not we agree on some arbitrary working commensurability
over the dimensions representing attributes (or some such measure as Fitch
and Margoliash’s “mutation distance”), in both space classes (I) and (II)
time is incommensurable with the other dimensions. We cannot legitimately
combine time and the other forms of “distance” into any general distance
measure. This is in fact the strength of the time-oriented standpoint; the
time-sequence is potentially determinable and often practically inferable.
Thus, the only unequivocal dendrograms are topological cladograms which
are qualitative in nature, except that if we do have palaeontological evidence
of actual times of branchings these cladograms become unequivocally metric
as to the time axis only.
CLADISTICS AND CLASSIFICATION
Assuming that we have arrived by reasonable inference at a cladistic
tree, does this then at last lead us to an optimal classification? It certainly
does not lead us to an ordination of the objects at any time level (more
strictly, in any hyperplane normal to the time axis), unless we accept a
particular attribute-space as in case (II) just described. But we may. consider
the possibilities of effective classification without ordination.
* Edwards and Cavalli-Sforza (1964, Cavalli-Sforza and Edwards, 1967) specifically
define an isotropic “evolutionary space-time’ but this isotropicity is achieved by a
transformation which, as they say, “is a reflection of the genetic assumptions which
are being made, and these will be peculiar to each case”.
= L. A. S. JOHNSON oe
Often, by reference to the time sequence of branching, we may refer the
objects to a nested (hierarchical) system of sets. However, if we are working
wholly or partly below the level of effective genetic isolation, the tree is a
banyan, anastomosing in its lower levels at least. The sets will then inter-
sect in a complex manner and in this case there is certainly no unique
hierarchy. This is one reason why the “biological” species concepts are
important in taxonomic theory and practice, however difficult to define they
may be in some groups of organisms (see Simpson, 1961, for an elaboration
of this matter, and Hull, 1964, 1965, for an illuminating philosophical
discussion). In these reticulate cases, Which are usually below and about the
“species” level in the conventional system, but may include somewhat higher
levels if allopolyploidy has oecurred, only subjective or arbitrary classifica-
tion is possible—there is no one “right answer”, though some classifications
are certainly worse than others. Numerical methods cannot alter this, though
they may aid in evaluation of data and obviate the possibility of grossly
subjective (that is, “unreasonable’’) judgements.
For many years I have worked with the large genus (or group of genera)
Eucalyptus, which presents a maze of anastomosing situations of this kind,
though not including alloploidy. It has long been clear to me that no single
“best” classification, phenetic or “biological”, is possible in such genera. It
would be a wasted effort to aim at one, and it is a fond hope indeed that
some numerical analysis or other is going to “solve” the classification problem.
Nevertheless, it is not wasted effort to attempt to understand the group or
to describe as well as we can the situations which exist within it, in terms
of phenetie variation, ecological relations, breeding behaviour, and so forth.
As Harlan Lewis (1957) has said, “it may be easier to determine why the
group is difficult than to decide the most appropriate taxonomic disposition”.
This is where the biosystematic way of thought has the advantage over
the formalism of the pure pheneticist. Biosystematics, even when not highly
experimental, is attempting to understand and describe an aspect of nature
in dynamic terms. This need by no means always, or even usually, be pushed
to the level of a detailed analysis of population dynamics; we can infer a
great deal from many other cases where detailed studies have been made.
We need to see the forest as well as the trees, and some of the trees as well
as the forest, but not always at the same time or place.
Formal taxonomy at the specific level cannot always aim at perfection,
though there are in fact many perfectly straightforward cases where no
dispute at all is likely to arise as to the correct grouping—nature often
does present us with comforting discreteness at a particular time level, hence
the not inconsiderable success of traditional systematics. In the difficult cases,
we can frequently improve the formal classification up to a certain point by
removing the grosser inconsistencies between formal treatment and our
biological understanding, although possibly replacing them with lesser
inconsistencies elsewhere. When we reach the stage where there is no gain
in consistency (this is itself not unequivocally measurable; see Hull, 1964),
we may as well describe the situation, and stop. Whatever theorists may
suppose, practising systematists are well aware that there are vast fields in
which much more effective work can be done before reaching the point where
imherent instability becomes limiting. Instabilities arising out of ignorance
or sheer wrong-headedness are, or were, common enough, but we need not
judge any subject by its incompetents. The improvement may sometimes be
made with the aid of numerical methods, but is often quite feasible without
them. Whether it is always worthwhile, by any method, is another question.
Ehrlich (1964, 1965) thinks not; he may be right, but this is a highly subjec-
38 THE QUEST FOR AN OPTIMAL TAXONOMY
tive judgement, like many pronouncements of the objectivists. Certainly some
more consideration of priorities in taxonomy, as suggested by Ehrlich, should
increase the efficiency of the subject as an aid to understanding. This is
quite apart from economic priorities, which are already well recognized.
Returning to a level above coenospecies and above alloploid formation,
where the cladograms are free of loops, the problems are again discussed by
Simpson (1961) and by Hull (1964, 1965). They are at two levels: phylo-
genetic interpretation and formal treatment. We shall here assume that our
interpretation is as good as we can get it—how, then, do we group, and
how do we rank?
Since lineages are continuous in time (up to the point of extinction),
classification along the time axis must be largely arbitrary, although the rate
of evolutionary change is by no means constant (see, e.g., Simpson, 1953)
and some authors (e.g., Takhtajan, 1953) have described evolutionary change
as chain-like rather than line-like. I cannot deal here with the difficulties
peculiar to the classification of allochronous forms. They are of much less
practical importance than those concerning contemporary organisms and may
be safely left to the palaeontologists. The text-book of Simpson (1961) and
the references given therein cover the subject adequately. Phyletic numerical
analysis may doubtless sometimes prove helpful in this field for the elucida-
tion of phylogeny and the quantitative study of anagenetic change, but any
purely phenetic analysis across different time levels is inappropriate for
general classification. Tuomikoski (1967) gives a useful discussion of palzon-
tological classification over all time levels but appears not to perceive the
virtual impossibility of a satisfactory solution.
Amongst organisms at any one time level, it may at first seem obvious
that, if we have deduced the topological structure of a divaricating tree and
expressed it as a cladogram, we then have a hierarchy laid out. before us.
The nested groupings would then present no problem though assignment of
rank would seem to have no non-arbitrary objective basis.
Unfortunately the situation is not so simple. Hull (1964) gives an admir-
able discussion of the difficulty of fitting the traditional Linnaean hierarchy
(indeed any hierarchy) to the phylogenetic background, and the reasons why
Simpson’s criterion of consistency can be only imperfectly applied. There are
two main sources of difficulty:
(1) The definiteness of a topological cladogram depends not only on our
knowledge, which may be quite inadequate to indicate the exact sequence of
branching, but also on the taxonomic wnit of the cladogram. Any questions
of monophyly will depend upon the unit adopted and here we encounter
some circularity in the definition of categories, if monophyly is taken as a
criterion (as it is by Simpson, 1961, and many other phyletic taxonomists).
A stem of a cladogram may be simple if we define it as being of generic cross-
section, as it were, at all points, but if we dissect it into species strands we
may find quite a network.
(2) Providing our representation of time and populations is sufficiently
coarse-grained, we may be able to say that species diverged at such-and-such
a level but it is not so easy with higher categories. This is fundamentally
because we insist on recognizing these categories by the “level of organiza-
tion” or “broad adaptive patterns” of the organisms we refer to them, as
well as by inferred monophyly (e.g., Simpson, 1953, 1961). This familiar
double function of the taxonomic categories was neatly expressed by Huxley
(1958) when he coined the terms grade and clade, now widely adopted. The
corresponding terms (due to Rensch, 1960) for processes of differentiation
2 L. A. S. JOHNSON 39
are anagenesis (evolutionary “advance”) and cladogenesis (evolutionary
separation), with the addition of stasigenesis (evolutionary stasis).
To illustrate this double requirement, the familiar example will serve:
The reptiles, mammals, and birds represent three grades, but cladistically
the last two are simply branches (clades, ignoring some parallelism in the
case of the mammals) of equal status to other branches still treated as
reptiles. In set-theoretic terms, the set: { all vertebrates whose ancestors
attained the reptilian level of organization } includes two (now) pretty
clearly defined non-intersecting proper subsets: {mammals} and { birds} ,
which we do not now call “reptiles”, together with the residue (the comple-
ment of the union of { mammals’ and { birds }) which we do call reptiles
but are jointly defined (within the original set) only by the fact that they
are not mammals or birds.
Ehrlich (1964) has said that “a system based on phylogenetic relation-
ships will not necessarily represent degree of phenetic differentiation”.
Restricting “phenetic differentiation” to what we regard as adaptationally
and practically important, this is quite true.
Now, in such cases the traditional hierarchy is not conformable with
the cladistic hierarchy, and the situation recurs throughout the living world.
It may be argued that the classification actually used is essentially phenetic.
So, in a sense, it is, but the choice of attributes used and the actual assign-
ment of individual taxa take evolutionary (but not only cladistic) considera-
tions very much into account.
Evolution, indeed, is not summed up merely in the topology of a phyletic
tree. Phenetic differences of many kinds (underlain by the genetic informa-
tion and the systems for its exchange, suppression, modification, and so forth)
are the very stuff on which selection works. As I have said earlier, because
we are part of the world in which selection operates and because what is a
practical matter for selection is often a practical matter for us, we are also
concerned with such phenetic characteristics, and a purely cladistic classifica-
tion will not meet our needs. Nor will a patristic system.
Thus we are presented with a dilemma: If we choose to classify purely
phenetically (this need not be hierarchic—overlapping clusters or simply
ordination are systematizations, though psychologically unsatisfying to most
of us) then we have no firm basis and are ignoring phylogeny, which does
interest us. If we choose to classify purely cladistically (assuming that we
have enough information) then we fail to display phenetic aspects of nature
which are highly important in the evolutionary process, to the organisms
themselves, and to ourselves.
The customary attempted solution of the evolutionary taxonomist is a
compromise. Its general rationale is given by Simpson (1961). It has been
followed, inevitably with varying degrees of imperfection, by many taxo-
nomists, sometimes avowedly (e.g., Johnson, 1959, pp. 76-77), sometimes
implicitly, and sometimes even when accompanied by disavowal.
Such a procedure is obviously not very logical and its defence may
present many problems. It is offensive to those who dislike compromise and
demand a clear codification of all procedures (e.g., Sokal and Camin, 1965).
But it serves our purpose very well, and we should think very hard before
we abandon it. For an anology we may look to language. Natural languages
are complex, not very logical systems, full of redundancies and overlaps.
Ordinary language will not do for mathematics or even for the discussion of
semantics itself, so we invent special symbolic languages and meta-languages.
Nevertheless, most of us have not yet found it necessary, feasible, or desirable
40 THE QUEST FOR AN OPTIMAL TAXONOMY
to discard common language even in scientific communication. With some
modifications and supplementation, it serves us better than any substitute
yet thought of, because it suits our psychology (which is far from simply
logical and linear) and because it links us with the knowledge of the past.
If, for similar reasons, we retain the broad framework of Linnaean
hierarchical taxonomy (though we may see fit to do away with some of its
more uselessly archaic features), we must face the fact that no optimal
taxonomy exists. We could discard Linnaean taxonomy for one of the
innumerable phenetic systems (there is no optimality there, as we have seen),
or we could use a purely cladistic system which could then (in theory) be
optimized, down to a certain level, as to the nesting of sets but not as to
ranking. As Hull (1964) has shown, this latter would result in a “pro-
hibitively complex and asymmetrical classification”, which would not meet
our requirements as we see them at present.
Recognizing and accepting the procedures which man has followed in
intuitive classification, Davidson (1967) suggests a radical departure from
the formal models employed by other taximetrists, whether phenetic or
phyletic. He recommends that mathematics should be used to construct a
cybernetic model congruent with the “classification program used by man
since time immemorial”. His efforts will be worth watching.
One of the logicians’ objections to the usual formal hierarchy has been
the recognition of categories which may contain only one member of next
lower rank, for example, unispecific (“monotypic”) genera. This objection
arises from a particular set-theoretic treatment under which the existence
of such categories leads to what has been called Gregg’s Paradox. This
paradox, like most others, is a consequence of the logical system adopted.
Under the more reasonable method of definition of taxon-names (in the sense
of Icgic) used in a logical analysis of the Linnaean hierarchy by Buck and
Hull (1966), the paradox vanishes. We can thus rest easy with a long-
established feature of systematics which is certainly meaningful to most
biologists.
The continual “improvement” of classifications (on whatever basis) leads
at present to some instability in names, which certain biologists and others
appear to find most vexatious (this is quite apart from formal changes due
to newly discovered nomenclatural priorities, and so forth). Systematists
themselves often do not mind these changes, at least in groups with which
they are familiar, since they feel that the new nomenclature expresses the
“cleaning-up” which has been achieved. Nevertheless, the instabilities do cause
confusion. Various authors have suggested that there should be a fixed
nomenclature, perhaps using single-word names for species, separate from
classification. Some (e.g., Michener, 1963; Sokal and Camin, 1965) sugggest
the use of code numbers. Hull (1966) proposes the adoption of Michener’s
fixed numbers supplemented by a system of “phylogenetic numericlature”
with adjustable “positional numbers” to indicate current views on phylo-
genetic arrangement. Perhaps some such system will eventually come, but
at present it seems premature. The mnemonic value of words, and of
binomials in particular, remains, despite challenges, and a great deal of
information in the literature would become difficult of access if the present
system were discarded. Similar views are expressed by Randal and Scott
(1967), who point out the usefulness of natural-language input to data-
processing machines, and also the greater detectability of errors by the eye
when words are used. Parkes (1967) gives a similar qualified defence of
traditional nomenclature, but we retain it chiefly, I think, fawte de mieuz.
L. A. S. JOHNSON 41
CONCLUSION
So, neither phyletics nor phenetics will lead us to the optimal classifica-
tion, the crock of gold, because it is not there. None the less, the elucidation
of phylogeny can still proceed, and our unperfectable classifications can still
be improved by reducing inconsistency until uncertainty or instability renders
further change unprofitable. Above all, taxonomists can set themselves the
task, as more and more of them do, of describing the situations they perceive,
giving reasons for their judgements, and suggesting lines of inquiry. The
investigation of phenetic and “biological” characteristics of organisms and
of their evolution is the scientific part of taxonomy. Classification itself
remains an art, but a disciplined and respectable art whose aim is to serve
science, not to express the personality of its exponents.
The classificatory component of taxonomy cannot itself be made into a
science by ill-founded philosophy or essentially arbitrary numerical pro-
cedures and taxonomists would be well advised to treat “philosophical”
pheneticism as unjustified in its claims and unproductive in practice. They
will surely find numerical techniques useful, particularly on the big problems,
and especially if the taximetrists can free themselves of their pheneticist
dogmas. Taxonomists may remember, however, that many other techniques
and tools, new and old, are at their disposal, including the subtlety of the
human intellect and its power of perceiving Gestalt and of bringing informa-
tion and theoretical reasoning of all kinds to bear on a problem.
If systematics is made into a sterile exercise, a purely pragmatic service,
or a playground for technicians, I would advise intelligent young biologists
to steer well clear of it. On the other hand it can be, as J have found it,
a stimulating and rewarding (though inherently inexact) branch of science,
with its dash of art, like most other stimulating branches of science, including
mathematics itself. To conclude with the words of that wise and wide-ranging
taxonomist, Lincoln Constance (1964), taxonomy in its widest sense, which
goes .far beyond the formulation of classifications, remains “an unending
synthesis”.
Acknowledgements
My colleague Dr. Barbara Briggs, by her helpful criticism, has prevented
this journey through the mists of theory and philosophy from being even
more disordered and discursive than it is; I am most grateful to her. My
wife has patiently typed illegible and tortuous manuscript, and borne with
long discussions; I am equally grateful to her. I wish to thank also (and
to ask tolerance for the cut-and-thrust of debate from) my friends Dr. W. T.
(Bill) Williams and Dr. Paul Ehrlich whose stimulating and provocative,
but also constructive, words and works have partly brought about the present
assessment of trends in taxonomy, with much of which IT naturally do not
expect them to agree!
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9-005
THE RHAPHIDOPHORIDAE (ORTHOPTERA) OF AUSTRALIA
PART 7%. PALLIDOTETTIX, A NEW GENUS FROM THE
NULLARBOR PLAIN, SOUTH-WESTERN AUSTRALIA
Aouta M. Ricwarps
Department of Zoology, University of New South Wales, Sydney
[Read 27th March, 1968]
Synopsis
A new genus Pallidotettix is erected, and the new species Pallidotettia
nullarborensis, nu. sp. is described from limestone caves on the Nullarbor Plain in
south-western Australia.
INTRODUCTION
The Nullarbor Plain is an area of Tertiary limestone about 75,000 square
miles in extent, situated in south-western Australia. It extends from Ooldea
and Colona on the east to Balladonia and Point Culver on the west, a
distance of about 500 miles. To the north it is bounded by the Great Victoria
Desert and to the south by the Great Australian Bight.
From 1957 onwards, a number of speleological expeditions have explored
many of the caves which occur throughout the southern part of the Plain,
and most of the material in this paper is based on the Rhaphidophoridae
collected on these trips. These insects have been collected from at least 24
of the known caves, and observed in another three caves. Their distribution
ranges from White Wells Cave, (N 14)* in the far east, to Gecko Cave (N 51)
at the farthest western extremity of the limestone and as far north as Lynch
Cave near the Transcontinental Railway (Text-fig. 1). Examination of these
specimens has shown them to belong to a single species which is described
here and placed in the new genus Pallidotettix, n.g. as Pallidotettiz
nullarborensis, n. sp. The species appears to be confined to the limits of. the
Nullarbor limestone.
P. nullarborensis is the palest Australian rhaphidophorid so far
examined by the author. Sexual dimorphism is not strongly developed. It
is absent from body size, antennae and number of linear spines on the hind
femora. It is present in leg length, although not as pronounced as in some
species of other genera. P. nullarborensis is one of the larger of the Aus-
tralian Rhaphidophoridae. An adult male may reach a length of up to
17 cm. from the tip of its antennae to its hind tarsi. It is of comparable
size to Novotettix naracoortensis Richards, but females are somewhat larger.
Pallidotettia does not show any close affinities with other Australian
rhaphidophorid genera so far studied. The structure of the external genital
plates in both sexes is quite distinctive.
Genus PALLIDOTETTIX, Nn. g.
Body sparsely clothed with short setae. Legs long and slender. Antennae
very long and tapering, almost touching at their bases; scape about three
times as large as pedicel, which is narrower than scape, but broader than
*The Nullarbor caves and dolines have been indexed by the Australian Speleo-
logical Federation, and the numbers are prefixed by the letter N. This system is
used throughout this paper. -
PROCEEDINGS OF THE LINNEAN Society or NEw SoutTH WALES, VoL. 93, Part 1
= AOLA M. RICHARDS 4G
other segments; from fourth segment onwards segments subequal in length,
although steadily decreasing in size; all segments thickly clothed with short
setae. A single anterior, median ocellus only. Fastigium rising very abruptly,
convex, grooved medianly and longitudinally. Fore coxae unarmed. All
femora sulcate ventrally. Apical spines on femora, tibiae, first and second
Immarna
Lake
aurice
M
wy
99 Ze
7,86
Location and index number of selected
Margin of Pleistocene and recent dunes
Chain of depressions along former
Margin of Tertiary limestone with other
watercourse
4 Koonalda
amen = Coastal cliffs (present and emerged)
Z
AO
Scale of Miles
d
~
N 5
8 oOo
w (Ss
& 8
~ a)
X
Rawlinna
Cocklebiddy
"156
Nullarbor Limestone, and the locations of the caves from which Pallidotettix
nullarborensis has been collected.
48 RHAPHIDOPHORIDAE (ORTHOPTERA) PART 7. PALLIDOTETTIX, A NEW GENUS
proximal segments of hind tarsi constant in number. Fore femur bears two
apical spines beneath, one prolateral and one retrolateral; fore tibiae bears
four apical spines, one above and one beneath both prolaterally and retro-
laterally; fore tarsus unarmed. Middle femur bears two apical spines
beneath, one prolateral and one retrolateral; middle tibia bears four apical
spines, one above and one beneath, both prolaterally and retrolaterally ;
middle tarsus unarmed. Hind femur bears two apical spines beneath, one
prolateral and one retrolateral; hind tibia bears a pair of long apical spurs
above, a pair of subapical spines above, a pair of shorter apical spurs
beneath and a pair of subapical spines beneath, one from each pair being
prolateral and the other retrolateral; two proximal segments of hind tarsus
each bears two apical spines above, one prolateral and one retrolateral ;
other two segments unarmed. Subgenital plate of female trilobed distally
and slightly keeled. Subgenital plate of male wider than long, distal
margin emarginate and medianly produced into a small lobe.
Type species for the genus: PALLIDOTETTIX NULLARBORENSIS, N. Sp.
PALLIDOTETTIX NULLARBORENSIS, Nl. Sp.
(Text-fig. 2, Figs 1-6)
Colour.—Basic colour ochreous, with pronotum, mesonotum, metanotum
and abdominal terga irregularly mottled with light brown; fore and middle
femora and tibiae of all legs ochreous, hind femora ochreous with transverse
light brown bands; tarsi ochreous; antennae ochreous; ovipositor light
reddish-brown.
Body.—tLength up to 16 mm. Body sparsely clothed with setae.
Ovipositor 0-8 length of body; ventral valves armed distally with six teeth,
gradually decreasing in size towards apex (Fig. 1). Antennae broken.
Fastigium longer than high with base touching scapes of antennae. Maxillary
palps with third and fourth segments subequal in length.
Antennae.—As in generic description. Third segment on dorsal aspect
2-25 as long as pedicel in female, and 1:75 as long in male; on ventral
aspect 1:25 as long as pedicel in female, 1-1 as long in male. Sexual
dimorphism absent. No spines on flagella of male or female.
Legs.—¥ore and middle legs subequal in length, with hind leg 1-5 length
of fore and middle legs. Sexual dimorphism slightly developed, fore, middle,
and hind legs of female being 0-9 as long as male. Hind femora, all tibiae
and proximal two segments of hind tarsi armed with variable numbers of
linear spines (Table 1). No spines occur on fore or middle femora and tarsi.
Apical spines constant in number as in generic description. Ratio of length
of legs to length of body: Fore leg, male 3°3:1; female 25:1. Middle leg,
male 3-1:1; female 2-4:1. Hind leg, male 4-8:1; female 3-7: 1.
Genitalia. Frmate: Suranal plate, Fig. 2 (SAP), concave laterally,
distal margin emarginate and clothed with setae; rest of plate sparsely
clothed with setae. Subgenital plate, Fig. 3 (SGP), 2-8 wider than long,
distal margin trilobed, two lateral lobes rounded and thickened along
margins, median lobe shorter and smaller than two lateral lobes and forming
a keel two thirds length of plate; whole plate almost glabrous. Mate:
Suranal plate, Fig. 4 (SPL), coneave laterally, distal margin slightly
emarginate with two latero-distal lobes; distal margin clothed with short
Setae, whole plate thickly clothed with setae, lateral lobes thickly clothed
with setae on ventral surface of plate (Figs 5, 6). Subgenital plate, Fig. 5
(Ff), 8-8 wider than long, convex laterally; distal margin slightly emarginate,
and medianly produced into a small lobe; whole plate sparsely clothed with
AOLA M. RICHARDS 49
T<—T VIII
Text-figure 2—Pallidotettix nullarborensis, n. sp. 1, Distal portion of ovipositor
showing teeth on ventral valve; 2, Female genitalia, dorsal view; 3, Female genitalia,
ventral view; 4, Male genitalia, dorsal view; 5, Male genitalia, ventral view; 6, Male
genitalia, ventral view, subgenital plate removed to expose structures beneath.
INDEX TO TEXT-FIGURE 2
BC, basal segment of cercus; C, cercus; DV, dorsal valve; H, subgenital plate,
male; IA, intersegmental apodeme; MTIX, membrane of tergite IX; P, paramere;
PVIII, pleurite VIII; PD, pseudosternite; PM, perianal membrane; PN, penis;
S, stylus; S VII, S VIII, S1IX—sternite VII, VIII, IX; SAP, suranal plate, female;
SGP, subgenital plate, female; SPL, suranal plate, male; T VII, T VIII, TIX, T X—
tergite VII, VIII, IX, X; 1 VF, first valvifer; 2 VF, second valvifer; VV, ventral valve.
D
50 RHAPHIDOPHORIDAE (ORTHOPTERA) PART 7. PALLIDOTETTIX, A NEW GENUS
setae. Two styli, Fig. 5 (S), broad, conical, thickly clothed with short setae,
length of styli being 1:6 length of sternite IX (SIX). Parameres, Fig. 6
(P), elongate with rounded apex, subequal in width to length, thickly
clothed with long and short setae. Pseudosternite, Fig. 6 (PD), twice as
wide as long, with rounded apex. Penis, Fig. 6 (PN), two-lobed, each lobe
subequal in width to length. Paraprocts, Fig. 6 (PP), twice as long as
wide, partially hidden between suranal plate and parameres, clothed with
setae.
Distribution.—Limestone caves on Nullarbor Plain, stretching from
White Wells Cave, South Australia, to Gecko Cave, Western Australia.
Gecko Cave (N 51), 15 miles east of Mt. Ragged, south of Balladonia,
W.A. (type locality), coll. D. C. Lowry 27/6/65; Lynch Cave (N 60), two
miles south-east of Loongana, W.A., coll. D. C. Lowry 12/5/66, A. M.
Richards 3/2/68; Pannikin Plain Cave (N 49), W.A., coll. D. C. Lowry
27/12/65; Tommy Graham’s Cave (N56), near Madura Pass, W.A., coll.
D. C. Lowry 11/12/65; Horseshoe Cave (N59), W.A., coll. D. C. Lowry
TABLE 1]
Variability in number of linear spines on the legs of Pallidotettix nullarborensis, n.sp.
Number Standard :
Mean Specimens Deviation Range
L R L R L R L R
Fore tibia Pro. 4-0 a9 25 25 0-3 0-4 3- 5 3- 5
Inf. Retro. 4-0 3°9 25 25 0:2 0:3 3- 4 3- 4
Mid tibia Pro. 3:6 3°6 25 25 0-5 0-5 3- 4 3- 4
Inf. Retro. 3°6 Si 25 25 0-5. 0:5 3- 4 3- 4
Hind femur Pro: 113 12-7 25 22 3:2 3°7 6-17 7-17
Inf. Retro. 14:6 15-6 25 22 6°3 4-5 7-35 8-26
Hind tibia 1EXRO}, 40:9 41-4 25 21 6-8 6-0 33-60 31-57
Sup. Retro. 43-6 43-7 25 21 5-8 7-1 3458 33-65
Hind tarsus iro: Bol!) 3°8 25 20 Holl 1-0 2- 6 2- 6
1 Sup. Retro. 3:8 3:8 25 20 1-0 0-9 2— 6 2- 6
Hind tarsus 127K), Ivo al 1-2 25 20 0-5 0-4 O- 2 l- 2
2 Sup. Retro. 1-0 1-0 25 20 0-4 0:3 0- 2 0- 2
INDEX TO TABLE 1
L., left leg; Inf., inferior; Pro., prolateral; R., right leg; Retro., retrolateral;
Sup., superior. 5
7/5/66; Blowhole (N52), 32 miles south-west of Rawlinna, W.A., coll.
D. C. Lowry 1/12/65; Moonera Tank Cave (N53), W.A., coll. 1D (Of Lowry
14/5/66; Murra-el-elevyn Cave (N 47), W.A., coll. I. D. Wood 3/1/64, P. F.
Aitken 3/1/64, E. Hamilton-Smith 28/12/64 and 17/1/65, D. C. Lowry
27/4/66, A. M. Richards 31/1/68; Cocklebiddy Cave (N48), W.A., coll.
P. F. Aitken 3/1/65, D. C. Lowry 12/1/66, A. Baynes 24/3/67; Unnamed
Cave (N57), 20 miles from Madura Pass, W.A., coll. D. C. Lowry 11/12/65;
Unnamed Cave (N 140), W.A., coll. D. C. Lowry 14/9/66; Unnamed Cave
(N 149), W.A., coll. D. C. Lowry 21/9/66; Old Homestead Cave (N 83),
W.A., coll D. C. Lowry 2/9/66; Snake Pit Cave (1383), Mundrabilla Station,
W. JN. coll. A. Baynes 27/3/67; Walpet Cave (N 38), W. A., coll. P. F. Aitken
January, 1964; Joe’s Cave (N39), W.A., coll. P. F. Aitken January, 1964;
Weebubbie Cave GNIZ) EWA collie thy Aitken 3/1/60, G. S. Hunt 29/12/65:
Murrawijinie No. 1 Cave (N 7), Nullarbor Station, S.A., coll. A. M. Richards
25/1/68; Murrawijinie No. 2 Cave (N 8), Nullarbor Station, S-A.., coll; AyM:
AOLA M. RICHARDS 51
Richards 25/1/68; Murrawijinie No. 3 Cave (N9), Nullarbor Station, S.A.,
coll. C. Warner February, 1957, A. M. Richards 26/1/68; Koonalda Cave
(N 4), S.A., coll: P. F. Aitken 7/1/59: A. Gallus January, 1968; White Wells
Cave (N 14), S.A., coll. P. F. Aitken 12/1/60; Koomooloobookka Cave (N 6),
Koonalda, S.A. coll. C. Warner February, 1957; New Cave (N 11),:S.A., coll.
N. Mollet January, 1955; Caves, Nullarbor, S.A. coll. J. Madden January,
1957; Cave, Nullarbor, S.A., coll. K. Renwick January, 1957; Cave, Nullarbor,
coll. unknown January, 1952. Also observed by D. C. Lowry during 1966 in
Unnamed Cave (N 156) near Balladonia, W.A., in Unnamed Cave (N 157)
south-east of Batladonia, W.A.; and in Unnamed Cave (N 158) near Cockle-
biddy, W.A.
Types.—Holotype male, allotype female, two paratypes, one male and
one female, in National Insect Collection, C.S.I.R.O., Canberra. Two para-
types, one male and one female, in Western Australian Museum Collection,
Perth; paratype, one female, in South Australian Museum Collection,
Adelaide.
Material examined.—Fourteen adults, 120 nymphs.
Acknowledgments
I wish to thank all the people who have helped in the collection of
specimens from various caves across the Nullarbor Plain. In particular
I should like to thank Mr. D. C.. Lowry, Geological Survey, Perth and his
wife for sending me specimens from the western part of the Plain. I am
also indebted to Dr. W. P. Crowcroft, Director of the South Australian
Museum, Adelaide, for the loan of material for study. I am grateful to
Mr. J. N. Jennings, Australian National University, Canberra for permission
to reproduce his map of the Nullarbor Plain, and to Mr. D. C. Lowry for
plotting on it the exact location of the caves. Finally, I wish to thank
Mr. C. J. Wilkinson, Geography Department, University of New South
Wales, for drawing the map.
A TAXONOMIC REVIEW OF THE GENUS MIXOPHYES,
(ANURA, LEPTODACTYLIDAE)
I. R. STRAvGHAN
Department of Zoology, University College of Townsville,
Townsville, Queensland
(Plates 1-11)
[Read 27th March, 1968]
Synopsis
Two species of Mixophyes Giinther are described and the two subspecies already
defined are elevated to species. ‘
INTRODUCTION
Moore (1961, p. 165) suggested that Mixophyes from northern New South
Wales might fall into two species—one from mountain streams and the other
from large coastal rivers. Analysis of additional material of Mixophyes
collected by the author from northern New South Wales and southern Queens-
land shows that three easily distinguishable morphological types, each with
a distinct call, are recognisable. Also the geographically isolated Mixophyes
from northern Queensland is distinct in morphology and call from any of the
southern forms. These four species are described in this paper.
MATERIAL AND METHODS
All the material listed by locality (tabulated north to south) was collected
by the author from breeding congresses during summers from 1960 to 1967
inclusive. This material is lodged with the Queensland Museum, Brisbane.
All material in the Australian Museum (Sydney) collections was examined.
Specimens examined by Moore (op. cit.) in the American Museum of Natural
History were not seen. Synonymies include the original reference, reference
to Parker (1940) who gives complete synonymies to that date, and all
subsequent references. The descriptions follow the pattern suggested by
Moore (op. cit., p. 155). Length of inner metatarsal tubercle was measured
along its long axis, and length of first toe from its tip to its junction with
inner metatarsal tubercle. A key to all the known species of Mixophyes is
provided.
MIXOPHYES FASCIOLATUS Gtinther
(eal ay Taives, 11)
Mixophyes fasciolatus Giinther, 1864, p. 46, pl. 7, fig. 1. Mirophyes fasciolatus
Giinther, Slevin, 1955, p. 359 [part.]; Moore, 1961, p. 162, fig. 1 [part.].
Mixophyes fasciolatus fasciolatus Giinther, Parker, 1940, with complete
synonymy (delete Fletcher, 1892, p. 18).
Type locality: Clarence River, N.S.W. Topotypic material 3 dd, 1 9,
collected two miles east of Grafton, Clarence River, 17-I11-1963.
PROCEEDINGS OF THE LINNEAN Society or NEw SourH WALES, Vor. 93, Part 1
= I. R. STRAUGHAN 53
Diagnosis: Species of Mixophyes are distinguishable from all other
Australian frogs by pupil vertical when constricted; vomerine teeth in front
of choanae; round tongue, only 1/4 to 1/5 free behind; distinct tympanum ;
limbs with dark cross bars; and feet with well developed web.
Mixophyes fasciolatus is distinguished from its congeners by weh to distal
sub-articular tubercle on outer margin of third toe; three joints of fourth
toe free of web; inner metatarsal tubercle as long as first toe; dark cross bars
on limbs narrower than intervening light; lateral zone of dark spots dividing
dark dorsal and white ventral surfaces; outer metacarpal tubercle well
developed; tympanum oval, long axis tilted towards eye.
Description: Large (60-80 mm., snout-vent length) with long barred
legs. Dorsal surface smooth to finely granular; tan to pale grey marked with
darker blotches: characteristically a dark mid-dorsal band present, com-
mencing as interorbital Y or T continuing posteriorly with irregular margins.
Band may be broken into less regular blotches, in extreme cases leaving only
interorbital bar and oblique elongate patches scattered over remainder.
Junction of dark dorsal and pale ventral surfaces marked by series of irregular
dark lateral spots between arm and groin. Distinct black head stripe from
behind nostril, through eye, above tympanum, curving round its upper margin,
ending at level of lower margin and separated from it by width of stripe
(Fig. 1c). Triangular patch in front of nostril with base along upper lip
Text-fig. 1. General head profiles and hind feet of Mixophyes, spp., showing head
stripes, shape of snout, position of tympanum, extent of webbing, and size of inner
metatarsal shovel. a.—M. balbus, b—WM. iteratus, c—M. fasciolatus, d.—M. schevilli.
and apex at nostril. Dorsal surfaces of limbs with dark cross bands not as
wide as intervening pale. Bands widening on margins of limbs to form dark
and light saw tooth pattern, breaking up irregularly on posterior surface
of thigh producing rough marble pattern. Ventral surface, including limbs,
white, smooth except for area around vent. Base of triangular expansion of
dark limb bars visible from below. Chin and throat dusted with dark.
Tympanum distinct, oval, long axis tilted towards eye from vertical (Fig 1c).
Fingers unwebbed, in order of length 3 > 4=1> 2. Elongate well-developed
inner, and smaller well-developed oval, outer metacarpal tubercle. Webbing
between toes strong. First toe free of web to proximal sub-articular tubercle,
as is second on inner edge. On outer edge of second, fringe of web to dilated
tip. Web only to proximal sub-articular tubercle of third on inner edge and
54 A TAXONOMIC REVIEW OF THE GENUS MIXOPHYES
to distal on outer edge. Three joints of fourth free. Fifth webbed to tip, but
web reaching only to level of distal sub-articular tubercle at lowest point
between fourth and fifth (Fig. 1c). No outer metatarsal tubercle but well-
developed, shovelshaped, inner metatarsal tubercle as long as first toe.
Sub-articular tubercles oval moderately developed (Fig. 1c). Vomerine teeth
platelike, directed towards each other from anterior margin of choanae to
midline between choanae, nearly touching.
Material examined: Queensland: Mount Cooroy 2 dd; Jimna Range 5 dd;
Nanango 3 6d; Kumbia 2 dd; Yarraman State Forest 3 dd; Yarraman
7 63; Bunya Mountains 17 dd, 3 ¥; Blackbutt 4 dd; Maleny 12 dd, 7 8;
Upper Brisbane River 6 dd; Chevellum 4 33; Woodford 3 dd; D’Aguilar
2 55; Mount Mee 14 dd, 11 $2; Ravensbourne National Park 1 ¢; Mount
Glorious-Mount Nebo 107 dd, 42 92; Samford 5 dd; Withcott 4 dd; Lake
Manchester 7 3d, 2 22; Gold Creek Road 4 63; Ma Ma Creek 1 ?; Mount
Tambourine 18 dd, 3 °2; Cunningham’s Gap 21 dd, 4 $2; Nerang River 3 dd;
Spicer’s Gap 1 d; Beechmont 1 °; Springbrook 11 6d; Binna Burra 4 dd;
Christmas Creek 14 dd; Coomera Gorge 1 3; Currumbin Creek 7 dd, 1 2;
Queen Mary’s Falls National Park 14 dd, 3 2; Mount Lindsay 2 dd. New
South Wales: Chillingham 5 dd, 1 ?;- North Arm Tweed River near
Murwillumbah 31 dd, 12 22; Island in Tweed River 2 dd; Woodenbong 3 dd;
Tweed River, Mount Warning 6 33; Richmond River (4 miles north of Kyogle)
1 3; Ulong 2 35; Upper Clarence River 7 6d; Richmond River (south of
Kyogle) 4 563; Grafton 3 3d, 1 2; Point Lookout, New England National
Park 51 dd. Australian Museum: 6791 3, Clarence River (N.S.W.) ; 6794 4G,
Pine Mountain (Qd.) ; R.4247 3, Warrell Creek, Nambucea (N.S.W.; R.5090 4,
Richmond River (N.S.W.) ; R.5872 ¢, R.5873 3b, R.5876 3b, R.5877 3, Nambucca
River (N.S.W.); R.6265 ¢, R.6287 3, R.6289 3g, R.6291 3, Gurravembi,
Nambucca River (N.S.W.); R.6499 Juvenile, Avoca, via Gosford (N.S.W.) ;
R.7463 3, Dunoon, Richmond River (N.S.W.); R.8472, Juvenile, Avoca, via
Gosford (N.S.W.) ; R.8937 3, R.8939 3, Mount Tambourine (Qd) ; R.10456 4,
Dorrigo Scrub (N.S.W.); R.10506 ¢, Wyong (N.S.W.); R.12081, Juvenile,
Palmdale, Wyong (N.S.W.); B.12645 3, R.12646 3, R.12647 3, R.12649 d,
R.12650 3, R.12651 3, R.12653 3, R.12654 d, R.12655 3, R.12656 3, Lowana,
Dorrigo (N.S.W.) ; R.13543 3, Old Koreelah (N.S.W.) ; R. 15127 3, R.16945 2,
R.17698 3, Bunya Mountains (Qd.) ; R.16922, Juvenile, Mount Glorious (Qd.) ;
R.20498, Juvenile, Guineacor Caves, Wombeyan Caves (N.S.W.).
Distribution: Along and east of the Great Dividing Range from Bundaberg
(Qd.) in the north to Gosford and Wombeyan Caves (N.S.W.) in the south.
MIXOPHYES ITERATUS, Sp. NOV.
(PIR igs 2)
Mixophyes fasciolatus Gimther; Fletcher, 1892, p. 18; Slevin, 1955, p. 359
[part.]; Moore, 1961, p. 163 [part.].
Type locality: Tweed River, Mount Warning, N.S.W.
Holotype: Australian Museum Reg. No. R.25929, d, collected 23-XI1-1963.
Paratypes: 12 Australian Museum; 1d, 1 2° Queensland Museum, collected
same time and place as holotype.
Diagnosis: Distinguishable from frogs of other genera by the six features
listed for M. fasciolatus, and from its congeners by web to tip of first, third,
and fifth toes; two joints of fourth toe free of web; inner metatarsal tubercle
strongly developed but relatively short—half as long as first toe; dark cross
bars on limbs as wide as intervening light; tympanum almost round, long
axis vertical; skin very granular on back and legs; and pointed snout.
a I. R. STRAUGHAN 5b
Description: Extremely large for Australian frogs (80-115 mm. snout-
vent), strongly developed hind legs and webbed feet, resembling Rana.
Holotypes: Back finely granular, dark olive to black almost obscuring
typical Mixophyes dorsal patterning. Dark head stripe broad, almost same
width throughout except above tympanum, where it narrows to thin line on
edge of supra-tympanic fold (Fig. 1b). Broad lateral band of spots tending
to irregular mottling between arm and groin. Dark cross bars on dorsal
surface of limbs as wide as intervening olive, not expanded on margins. On
posterior surface of thighs, cross bars coalese forming uniform dark back-
ground with few distinct yellow spots of diameter approximately equal to
width of cross bars. Ventral surface smooth, white on belly limbs. Fine darker
dusting on chin. Tympanum distinct, almost round, long axis vertical. Sharp
supra-tympanic fold (Fig. 1b). Fingers without web, arranged in order of
length: 3 > 4>1 > 2. Inner metacarpal tubercle, oval, well developed. Outer
almost flat on palm. Nuptial pad on first finger only thin strip along inner
edge. Toes fully webbed, reaching tip of first, second, and third toes on outer
margins and tip of fifth. On inner edges of second and third to proximal and
distal sub-articular tubercles, respectively. Only two joints of fourth toe free
of web, narrow fringe to tip on outer margin (Fig. 1b). Inner metatarsal
tubercle without well-developed shovel edge, length equal to half of first toe
(measured from tip to its junction with tubercle). Sub-articular tubercles
elongate, flattened. No outer metatarsal tubercle. Vomerine teeth in slightly
oblique transverse series, almost meeting in midline, almost entirely in front
of choanae.
Snout-vent length 80-5 mm.
Variation: Three paratypes and other specimens examined vary little from
holotype. Females larger than males (> 100 mm.), skin with texture of coarse
sandpaper. Dorsal colour from pale olive to dark bottle green.
Material examined: Queensland: Kumbia 1 3; Bunya Mountains 3 3d, 12;
Cunningham’s Gap 5 dd; Queen Mary’s Falls National Park 2 dd, 2 99;
Mount Lindsay 2 dd, 1°. New South Wales: Upper Richmond River (6 miles
north of Kyogle), 7 dd, 1¢; Tweed River, Mount Warning, 4 dd, 2°. Australian
Museum: R.7493 °, R.7494 2, Dunoon, Richmond River (N.S.W.) ; R.7550 Z,
Dorrigo (N.S.W.) ; R.12308 °, Mullumbimby (N.S.W.) ; R.12642 3, R.12648 2,
R.12644 ?, Lowana, Dorrigo (N.S.W.); R.16762 °, Coolmangar, via Lismore
(N.S.W.) ; R.16946 o Mullumbimby (N.S.W.) ; R.19038 2, R.19039 2, R.19040 °,
R.19041 3, R.19042 3, Wallaby Creek, Urbenville (N.S.W.) ; R.25877 2°, Ourimah
( N.S.W.).
Distribution: Bunya Mountains and along the Queensland-New South
Wales border east of Stanthorpe, south to the Dorrigo Plateau, N.S.W.
MIXOPHYES BALBUS, Sp. Nov.
(edleciag, AMfease 3b)
Mixophyes fasciolatus Gtinther, Moore, 1961, p. 162.
Type locality: Point Lookout, New England National Park, N.S.W.,
between 4,250 and 4,750 feet altitude.
Holotype: 3 Australian Museum Reg. No. R.25922 collected 15-X-1965.
Paratypes: 10 53, 3 22 collected same time and place as holotype, by I. R.
Straughan and A. R. Main (Australian Museum and Queensland Museum).
Diagnosis: Distinguished from other Australian frogs by six features
listed for M. fasciolatus; and from other Mixophyes species by web extending
only to distal sub-articular tubercle of third toe on outer margin, three joints
56 A TAXONOMIC: REVIEW OF THE GENUS MIXOPHYES
of fourth toe free of web; inner metatarsal tubercle well-developed shovel,
equal in length to first toe; cross bars on dorsal surface of limbs narrow, not
distinct over whole surface, without distinct triangular bands on margins of
limbs; dorsal surface diffuses laterally to merge with white ventral, without
sharp change marked by narrow zone of dark dots; males with well-developed
nuptial pads on metacarpal, first and second fingers; and oval tympanum,
long axis directed obliquely towards eye.
Description: Large frogs (60-80 mm. snout-vent length) with strong limbs
poorly marked by dark cross bars.
Holotype: Dorsal surface yellowish grey (grey in alcohol) diffusing
gradually into white ventral. Lateral surface not marked with dark spots.
Dark markings of typical Mixzophyes pattern—interorbital T extending
posteriorly as broad mid-dorsal stripe of irregular outline; few scattered
irregular dark patches on remainder of back. Dark head stripe, bold between
nostril and eye, thin line above tympanum (Fig. la). Triangular patch in
front of nostril, not as dark as head stripe, with well marked edge. Bars on
dorsal surface of limbs narrower than intervening light, not distinct over
whole surface, broadening terminally, but not forming distinct triangles on
margins of limbs. Extremities of dark limb bars not visible from below.
Posterior surface of thigh diffusely speckled with dark. Ventral surface of
body and limbs white, hands and feet darker, chin dusted with darker.
Tympanum distinct, dorsal margin obscured in head stripe and supratympanic
fold, oval, long axis directed towards eye (Fig. la). Fingers without web,
stouter than in other species of Mixophyes, in order of length 3 > 4 >1=2.
Inner metacarpal tubercle elongate, strongly developed; outer oval and
equally developed. Dark horny nuptial pad covering dorsal surface of first
finger except for distal phalanx; separate round pad on inner surface of inner
metacarpus and tubercle; and thin strip dorsally on inner edge of second
finger. Toes webbed to: sub-articular tubercle of first, proximal sub-articular
tubercle of second and third on inner margins, tip and distal sub-articular
tubercle on outer margins of second and third respectively, and tip of fifth.
Three joints of fourth toe free of web, narrow fringe along outer edge to tip
(Fig. la). Inner metatarsal tubercle strongly shovel-shaped, basal length
approximately equal to length of first toe (measured from tip of toe to
junction with tubercle). No outer metatarsal tubercle. Sub-articular tubercles
variably developed. Vomerine teeth typical of Mixophyes—transverse plates
in front of choanae almost joining in midline, directed slightly backwards
towards midline.
Snout-vent length = 75:0 mm.
Variation: Webbing consistent on both sexes. Females without nuptial
pads, more slender fingers. First, second, and fourth fingers almost equal in
length, not always in same order as type (any order possible). Dorsal pattern
with similar variation to other species of Mixophyes—less regular dark
markings. Spots of dark in groin and behind arm in some females, not
marking a zone of sharp transition from dark dorsal to white ventral
colouring.
Material AUT New South Wales: Point Lookout, altitude 4,500 to
4,750 feet, 14 dd, 5 92, 16-X-1965; Point Lookout, ca. 4,250 feet, 27 dd (21
Sympatric with ie US GLOITEDS) 19-11-1966. Australian Museum: R.7479 Z,
Kurrajong Heights (N.S.W.) ; R.7567, Juvenile, Illawarra (N.S.W.) ; R.7587 3,
Burrawang (N.S.W.) ; R.8328 3, Mount Wilson (N.S.W.); R.8455 4, Moss
Vale District (N.S.W.); R.9218 3, Williams River, Dorrigo (N.S.W.) ;
R.10063.3, Blackheath (N.S.W.) ; R.12547 3, Mount Irvine (N.S.W.) ; R.12567
= I. R. STRAUGHAN BI
3, Mount Wilson (N.S.W.) ; R.12633 d, R.12648 3, R.12652 3, Lowana, Dorrigo
(N.S.W.) ; B.12788 , R.12789 2, Mount Irvine (N.S.W.; R.17086 3 .R.17087 2,
R.17131 2, Dorrigo (N.S.W.) ; R.17097 3, R.17581 3, Point Lookout, via Ebor
(N.S.W.) ; R.17671, Juvenile, Linden, Blue Mountains (N.S.W.) ; R.18555 2,
Barrington Tops (N.S.W.); R.19178 3, R.19179 3, R.19180 2, Royal National
Park, Sydney (N.S.W.); R.19257 3, 15 miles SE Moss Vale (N.S.W.);
R.19430 3, Mount Wilson (N.S.W.) ; R.24493 3, R.24494 do, R.24495 3, R.24496
3, R.24497 3, R.24498 S, Falconbridge (N.S.W.).
Distribution: Hast of the Great Dividing Range from the Dorrigo Plateau
south to Illawarra
MIXOPHYES SCHEVILLI Loveridge, new combination
(Vedle) toe, «Li iees, » 2)
Mixophyes fasciolatus schevilli Loveridge, 1933, p. 56; Parker, 1940, p. 15.
Type locality: Millaa Millaa; Lake Barrine, 4,000 feet Bellender Ker
Range, North Queensland.
Topotypic material: Millaa Millaa, 1 3, 2 ?2, collected 11-11-1963; Lake
Barrine, 1 ? collected 11-X1J-1964.
Diagnosis: Distinguished from other genera by six features listed for
M. fasciolatus and from other species in the genus by toes strongly webbed
with only two joints of fourth toe free of web, web to tip of third toe on outer
edge; basal length of inner metatarsal tubercle only approximately half length
of first toe; smooth skin, yellow-brown to red colour; cross bars on limbs
narrow, alternating with fine dark lines, passing completely round forelimb,
tibia, and foot; back of thigh with dark diffuse broad horizontal band formed
from coalesing. of cross bars, without pale spots; and oval tympanum, long
axis tilted towards eye from vertical.
Description: Large raniform frogs (60-90 mm. snout-vent). Dorsal surface
smooth, yellow-brown, tan, or darker brown with red tinge, back pattern
typical Mixophyes—dark interorbital T or Y continuing backwards as
irregular broad mid-dorsal stripe and irregular scattered blotches over
remainder. Dark head stripe with continuous black upper margin, broken on
lower margin between nostril and eye (Fig. 1d). Triangular patch in front
of nostril edged in black, otherwise paler. Dorsal surface of limbs with
distinct dark cross bars alternating with finer dark lines. Bars narrower
than intervening background, continuous across ventral surface round fore-
limb, tibia, and foot. On thigh, bars coalese and posteriorly form a diffuse
dark horizontal band, not marked by paler spots. A few scattered lateral dark
spots. Ventral surface of body and thighs white, smooth except immediately
adjacent to vent. Throat white, chin dusted with black. Tympanum oval,
long axis tilted from vertical towards eye (Fig. 1d). Fingers not. webbed,
in order of length 3 >4>1>2. Inner metacarpal tubercle elongate to
oval well developed; outer oval, barely raised from palm. Toes webbed to:
just beyond sub-articular tubercle of first, tip of second and third on outer
margins, proximal sub-articular tubercle and slightly beyond on inner margin
of second and third respectively, and tip of fifth. At most, two joints of
fourth toe free of web (Fig. 1d). Inner metatarsal tubercle shovel-shaped,
without well developed edge, length approximately half length of first toe
from tip of toe to junction with tubercle. Sub-articular tubercles oval,
flattened. No outer metatarsal tubercle. Vomerine teeth oblique transverse
plates directed from front margin of choanae to midline between choanae.
58 A TAXONOMIC REVIEW OF THE GENUS MIXOPHYES
Material examined: Queensland: Mount Finigan (Cooktown) larvae only ;
Black Mountain (20 miles north of Kuranda), 2 dd, 3 ¢%; 7 miles west of
Atherton, ca. 4,000 feet, 2 dd, 1 °, 23 Juveniles; Mount Hipipamee, 1 d, 1 ¢;
Lake Eacham, 1 d, 1 2; Lake Barrine, 1 ?; Malanda, 1 3; Millaa Millaa, 1 d,
2°22; Tchupala Falls, Palmerston National Park, 2 dd, 2 Juveniles. Australian
Museum: R.266 3, 20 miles inland of Cairns (Qd.) ; R.770 3, R.4693 Juvenile,
Cairns District (Qd.) ; R.17017 3, Dinner Creek, near Cairns (Qd.).
Distribution: Atherton Tablelands and coastal ranges of north Queens-
land from Mount Finigan (near Cooktown) south to the Johnstone River.
Key to all known species of MIxopHYES
1. Toes webbed to tip of third toe on outer margin; only two joints of fourth toe free
of web. Basal length of inner metatarsal tubercle half length of first toe
(measured from tip of toe to its junction with metatarsal tubercle) ........ 2.
Toes not webbed beyond distal sub-articular tubercle of third toe on outer margin;
three joints of fourth toe free of web, except for a fringe along outer margin.
Inner metatarsal tubercle ecqualjin’ leneth=to first) Toews... 4- aoe eee ee a.
2. Dark cross bars on limbs as broad as intervening light, coalescing on posterior
surface of thigh to form uniform dark background with scattered pale blotches
Fe HAN ie AMO eo oer AAD eg A eee ern CDR Ea dias MeN id oat eRe D105 Mixophyes iteratus.
Dark cross bars on limbs narrower than intervening light, coalescing on posterior
surface of thigh to form diffuse dark speckled, horizontal band ...............
Ee hich sam eS ein era aioe canes nell yh a Gi sihaisne clei s+ Jee vein) ae et MAC ODNYESSCILEULLt.
3. Dark cross bars on limbs sharp, well defined; widening on the margins into dark
triangles, the bases of which are obvious from below ..... Mixophyes fasciolatus.
Dark cross bars not sharply defined, with few irregular marginal expansions which
are motwvAsiblestromyibelowenc)-e aoa oe eee Mixophyes balbus.
DISCUSSION
Mixophyes fasciolatus Giimther as redefined here, is synonymous with
M. fasciolatus fasciolatus Gimther, defined by Loveridge (1933, p. 55) and
followed by Parker (1940, p. 18) who gives a complete synonymy, in which
only one reference does not now refer to this species, i.e., Fletcher (1892, p. 18)
referring to specimens with fully webbed toes from the Tweed River, N.S.W.
Slevin (1955) referred three specimens from Ulong, Richmond River, N.S.W.,
two with fully webbed toes, to M. fasciolatus. Moore (1961) found specimens
from Lowana, Mullumbimby, and Dunoon (all in northern N.S.W.) as well
as the specimens of Fletcher (1892) and Slevin (1955) were webbed to the
extent considered as diagnostic of M. fasciolatus schevilli Loveridge. As this
sub-species. was erected for northern Queensland forms and M, fasciolatus for
southern Queensland-northern N.S.W. forms, Moore (op. cit.) believed that,
because the two forms occurred in sympatry, sub-species could not be
recognised, He suggested that two species might be involved: a highland
rapids species with “mountain brook” tadpoles and a coastal stream species
with more extensive webbing and probably with unspecialised larvae. These
“fully webbed” specimens have been transferred to the new species M. iteratus.
Specimens of WM. balbus recorded in the literature under M. fasciolatus are
listed under Australian Museum numbers for M. balbus and may be cross
referenced to Moore’s list (p. 164). Moore (op. cit., p. 163) refers tadpoles,
found at 5,000 feet at Point Lookout, N.S.W. to M. fasciolatus, but as this
is the type locality for M. balbus, and M. fasciolatus is known only at lower
altitudes, this reference is more likely to be to M. balbus.
M., fasciolatus is sympatric with WM. iteratus and M. balbus in different
parts of its range and exhibits no integrading with -either. M. balbus and
M. iteratus occur sympatricly at several localities where no intergrades have
Proc. Linn. Soc. N.S.W., Vol. 98, Part 1 PLATE
Fig. 1. Mixophyes fasciolatus.
Fig. 2. Mixophyes iteratus.
fy ir i
ap
Proc. Linn. Soc. N.S.W., Vol. 938, Part J PLATE IT
Fig. 1. Mizrophyes balbus.
Fig. 2. Mirophyes schevilli.
S Ag
0 1
2 ~ 5
j
+ f Hy
2 ”
3]
y a
‘
5 a
j
ie ,
: are
iy
a a .
~
>see aN Ot eos cee i)
si hepa orst Ye
, 4
Bs
t
¢
x t
~
*%,
ie et ’ ;
= I. R. STRAUGHAN 59
been collected. M. schevilli is restricted in distribution and is geographically
isolated, but exhibits greater differences in morphology from the other species
of Mixophyes, than is found between these species. Straughan (1966) showed
that each of these species has a distinct mating call and found no intergrading
in areas of sympatry of the three southern species. Also the call of M. schevilli
reflects the degree of difference from the southern species shown by morphology.
Parker (1940, p. 15) gives a complete synonymy for M. schevilli.
References
FietcHer, J. J., 1892—Contributions to a more exact knowledge of the geographical
distribution of Australian Batrachia. No. 3. Proc. Linn. Soc. N.S.W., ser. 2, 7: 7-19.
Gutnruer, A., 1864.—Third contribution to our knowledge of batrachians from Australia.
Proc. zool. Soc. Lond.: 46-49.
LoveringE, A., 1933.—A new genus and three new species of crinine frogs from Australia.
Occ. Pap. Boston Soc. nat. Hist., 8: 89-94.
Mooreg, J. A. 1961.—The frogs of eastern New South Wales. Bull. Am. Mus. nat. Hist.,
121 (3): 149-386.
Parker, H. W., 1940.—The Australasian frogs of the family Leptodactylidae. Novit.
Zool., 42: 1-106.
SLEVIN, J. R., 1955.—Notes on Australian amphibians. Proc. Calif. Acad. Sci., 28: 355-392.
STRAUGHAN, if R., 1966—An analysis of species recognition and species isolation in
certain Queensland frogs. Unpublished doctoral thesis, University of Queensland.
EXPLANATIONS OF PLATES I-II
PLATE I
Fig. 1. Mixophyes fasciolatus (natural size). Dark dorsal and limb patterns obscured
by the overall darkness of this highly hydrated specimen. Dark sawtooth pattern on
margins of limbs and lateral series of dark spots obvious.
Fig. 2. Mixophyes iteratus, sp. nov. (4 x natural size). 9 Paratype Qd. Mus. showing
fully webbed feet, wide cross bars on limbs and granular skin.
PLATE II
Fig. 1. Mixophyes balbus, sp. nov. (natural size). Holotype Aust. Mus. Reg. No.
R.25922 showing two joints of third toe free of web; diffuse limb bars with irregular
expansions at margins, absence of lateral series of dark spots, and nuptial pads on first
and second toes.
Fig. 2. Mixophyes schevilli (natural size). Showing irregular cross bars narrowing
to and alternating with thin dark lines.
THE SECRETORY CAPACITY OF THE STOMACH OF THE WOMBAT
(VOMBATUS HIRSUTUS) AND THE CARDIOGASTRIC GLAND
G. W. Mitton,* D. J. Hineson; and E. P. Grorent
(Communicated by Dr. Mervyn Griffiths)
[Read 27th March, 1968]
Synopsis
The secretory capacity of the stomach of the wombat (Vombatus hirsutus), and of
the cardiogastric gland of this animal has been studied.
It was found that the secretory power of this stomach resembles that of man and
animals commonly used in gastric research. The concentrations of ions in gastric juice
generally fitted the Hollander two-component theory of gastric secretion. A close
correlation between the concentration of pepsin and of K* was demonstrated. The
electrophoretic pattern of the gastric juice of the wombat resembled that obtained from
the gastric juice of man. The maximum secretory capacity of the stomach of this
animal was lower than that of man. It was found that a considerable increase in the
gastric output could be obtained by augmenting the effects of histamine stimulation
by injections of insulin. :
INTRODUCTION
In three mammals, the wombat (Vombatus hirsutus), the koala “bear”
(Phascolarctos cinereus) and the North American beaver (Castor canadensis)
there exists on the lesser curvature of the stomach a highly specialised area
of secreting cells, known as the cardiogastric gland (Home, 1808; Johnstone,
1898; Mackenzie, 1918; Milton, 1962). The anatomy of this gland has been
studied in the beaver (Smith et al., 1911; Nasset, 1953) and in detail in the
wombat (Hingson and Milton, 1967). In the latter animal the gland consists
of about 20 infoldings of the gastric epithelium with the muscularis mucosae
and submucosa. Hach of the gland pits opens into the lumen of the stomach
through a separate ostium. The pits are lined with thickened gastric epithelium
which contains innumerable gastric glands. The cells in these gastric glands
are of the usual type found in the lining of a carnivorous mammalian stomach.
The chief and parietal cells are particularly prolific and the gland tubules
appear to possess a good blood supply from the left gastric artery. A separate
branch of the vagus nerve enters the gland from the parietal surface. The
remainder of the stomach in the wombat has a lining similar to that of other
mammals, the antrum is well defined and consists of non-acid secreting
epithelium. There is also a small band of non-acid secreting epithelium close
to the oesophageal opening. The body of the stomach is lined with glands
similar to other animals. The glands here consist of surface mucus cells,
mucus cells at the neck of the gland, chief and parietal cells and an occasional
argentaffin cell.
Kach of the animals possessing a cardiogastric gland is a herbivore. The
precise diet of the wombat is not fully known, but it probably consists of
roots and grasses that are found near its burrow. Both the wombat and the
* Department of Surgery, University of Sydney, Sydney.
y Research Student, Johns Hopkins University, Baltimore, U.S.A.
t School of Physics, University of New South Wales, Sydney.
PROCEEDINGS OF THE LINNEAN Society or New SoutH WALES, VOL. 93, Part 1
G. W. MILTON, D. J HINGSON AND E. P GHORGE 61
koala are large marsupial animals and are therefore widely separated in
evolution from contemporary mammals such as the beaver.
The object of the present paper is to report a study of the secretory
function of the wombat stomach and of the cardiogastric gland. We have
also investigated the ionic relationships in the gastric juice of this animal
to ascertain if they are in agreement with the Hollander two-component
theory of gastric secretion (Hollander, 1932).
MATERIAL AND. METHODS
Nine adult healthy wombats weighing between 13-25 kg. were used in
these experiments. The animals were fasted for 18—24 hours and anaesthetised
with nitrous oxide and oxygen mixture administered through a face mask.
As soon as anaesthesia had been induced a fine intravenous catheter was
inserted into one femoral vein and passed into the inferior vena cava. The
catheter was attached to a threejway tap and through it a connection made
to a syringe containing dilute pentothal sodium (1 gm./60 ml.). Anaesthesia
was maintained by occasional injection of 1—3 ml. of this solution. Throughout
all experiments the animal was given a continuous infusion of histamine acid
phosphate (2:2 pg. base/kg. body wt./min.) administered by a slow infusion
pump. In several experiments once a steady baseline of secretion had been
established booster doses of histamine were administered (0-5 mgm./kg.)
subcutaneously. After some hours of histamine the animals were given a
subcutaneous injection of soluble insulin (50 units).
The experiments were divided into two groups. In Group 1 the secretion
from the whole stomach excluding the antrum was studied. An intragastric
tube was passed through the mouth and its position checked by opening the
abdomen. A tight ligature was then passed around the antral region just
distal to the end of the tube, thereby excluding the antral secretion from the
rest of the stomach. In Group 2 the stomach was opened along the anterior
surface and a suction tube passed through the mouth and fixed over the
openings of the cardiogastric gland by sewing the tip of the tube to the lesser
curve just beyond the gland. The gland area was then excluded from the
remainder of the stomach by sewing together the anterior and posterior gastric
surfaces across the tube with an atraumatic stitch. In this way a pouch was
made which consisted largely of the cardiogastric gland. A second tube was
placed in the remainder of the stomach through the gastrotomy opening and
brought out through the anterior abdominal wall. The tubes used to aspirate
the gland area and the stomach were double lumened, the smaller lumen being
an air inflow to prevent excessive suction. When the tubes were in place the
caudal end of the table was raised eight inches to prevent the saliva from
trickling into the stomach and to facilitate aspiration.
At. the conclusion of the experiment the position of the tubes was checked
and also the patency of the septum between the cardiogastric gland and the
body of the stomach.
The samples of gastric juice were collected under ice and the collecting
cylinder changed half-hourly. After insulin injections the collecting cylinder
had to be changed more frequently (10-15 minutes). Each specimen was
tested for the following: (1) Acidity by titration against NaOH (N/50) to
pH 7-4, using a glass electrode pH meter coupled to an automatic titrator
(Radiometer). (2) Na* and K* by flame photometry (Perkin Elmer, external
standards). (3) Cl by potentiometric titration using the automatic titrator
(Lehmann, 1939; Muller, 1942). (4) Pepsin using the haemoglobin substrate
method (Harrison, 1964). (5) Total osmolarity was measured in seven
samples of high concentration using a freezing point osmometer. (6) In one
62 STOMACH CAPACITY AND CARDIOGASTRIC GLAND OF THE WOMBAT
animal the stomach was irrigated with NaHCO: by the technique of Piper
et al. (1963) and the electrophoretic pattern of the gastric juice estimated
by Piper (1966).
Samples of blood were taken from five animals and the total osmolarity,
Na*, K*, and Cl concentration of the plasma were measured by the above
techniques.
The figures were analysed in several ways. The interrelationships of ions,
volume, pepsin for both concentration and output were determined with the
assistance of the digital computer, sttuiac. The correlation coefficients between
pepsin and H’, K’, and Cl were obtained. In addition observations were made
on the response to histamine, and histamine augmented with subcutaneous.
histamine and subcutaneous insulin.
RESULTS
The characteristics of the serum of the wombat are shown in Table 1,
together with the total osmolarity of gastric juice secreted at maximal acid
concentration. It can be seen that the osmolarity and concentration of ions
in the serum of these animals is roughly the same as that in other animals
more frequently used in gastric research. The osmolarity of human, dog, and
cat plasma ranges from 300 to 330 m. osmoles/litre (Houssay, 1955; Spector,
1956). :
TABLE |
Serum osmolarity and the osmolarity of concentrated gastric
juice in milli osmoles|litre, and the concentration of Nat, K+
and Ol- (mEgq|l.) from the serum of the wombat, +S.D.
The figure in brackets is the number of specimens analysed
Total osmolarity serum Pie we 286-2+20-8 (5)
Tons (serum) : ;
ING 5 ic aI she is a 134-8+ 3:3 (5)
K ae =e ae is ee 4-44 0-35 (5)
ON ior a ih a he se 96-8+10-1 (5)
Gastric juice (whole stomach) Ais 281-4+ 33-2 (7)
The electrophoretic pattern of the gastric juice showed a main band that
moved towards the anode and had the electrophoretic mobility of albumin.
In addition there was a more rapidly moving anodal band (Band 6) and
several bands of intermediate mobility. The latter are almost entirely muco-
polysaccharides. The pattern resembles that found in human gastric juice
(Piper, 1966; Piper et al., 1963).
The response of the whole stomach (Group 1) and the isolated cardio-
gastric gland (Group 2) to histamine infusion was similar to other animals.
After the start of the infusion the volume and acidity of gastric juice increased
until after about one hour a peak was reached (Fig. 1). There was a tendency
after the establishment of a peak for the secretion rate to diminish, while the
acidity remained at a plateau. Supplementary subcutaneous injections of
histamine caused a temporary increase in gastric output, but no increase in the
concentration of acid. The injection of insulin was followed by a considerable
Increase in both the volume and the acidity of the gastric juice. The volume
of gastric juice and the output of acid taken over all experiments were closely
related (1 = 0-80).
The pepsin concentration remained at a high and fluctuating level for
some hours and then declined. The correlation of pepsin on volume was poor
(r = 0-29), neither the injection of histamine nor of insulin affected the decline
of peptic concentration. A typical result is shown in Fig. 2. The concentra-
G. W. MILTON, D. J HINGSON AND E. P GEORGE 63
tion of K* closely followed the concentration of pepsin in all the fluctuations
(x = 0-63).
The concentration of all the ions measured in all experiments followed
the usual relationships. Na* concentration fell as the acidity rose, and Cl and
K* followed each other closely (Table 2).
12
10
VOLUME mis,
Or
a)
fo)
HOURS
Fig. 1. Volume of gastric juice secreted from the whole stomach preparation
described in the text. Histamine stimulation was commenced at 0 time. Note the rise
to a peak after 1 hour and the decline despite continuing stimulation. Booster doses:
of histamine produced some increase in output, but a considerable increase in output
followed the administration of insulin. H—histamine, I—“insulin.
When comparing the maximum secretory capacity of the cardiogastric
gland (Group 1) and the whole stomach including the cardiogastric gland
(Group 2) it is necessary to take into account the effectiveness of the septum
TABLE 2
Correlation coefficients (r) for acidity (H) on Cl, Na, K and also for
pepsin (P) on H, Na, Cl and K
The concentrations of all ions were in mEq/l. and pepsin in
10? units/1.
L.C. Tons r Pepsin (P) P
H Vs Cl ore as 0-78 1D \WWiseisl aes —0:21
ESViss Nae: rR —0-83 12 Wey Cll 5 0-05
H Vs K Ae BE —0:27 PR) Vs Na ae —0-05
PVs K a 0-63
separating the gland from the body of the stomach. In one animal this septum
was not adequate to exclude leakage from the body of the stomach into the
cardiogastric gland area; the animal was excluded when the comparisons
were made. Table 3 shows the maximum concentration of ions excreted by
«64 STOMACH CAPACITY AND CARDIOGASTRIC GLAND OF THE WOMBAT
four isolated cardiogastric gland areas and the maximum concentration from
the whole stomach. There was no significant difference between any of the
figures obtained for H*, Cl, Na’, K° and pepsin (Table 3).
The Hollander (19382) two-component theory of gastric secretion is based
on the assumption that gastric juice is made up of two isotonic solutions.
700
600
500
PEPSIN x10° units /ml
400
K mEq/|
300
200
HOURS
Fig. 2. Concentration of pepsin (Units 108/ml.) and K* (mEq/Il.) from the gastric
juice obtained from the animal shown in Fig. 1. The concentrations of pepsin and acid
show similar fluctuations.
If this assumption is valid for the wombat there will be a linear relation
between the concentrations x, y of different ions, taken two at a time, of
the form— y=at bx (1)
where the constant b is given by
xy being the co-variance
xx being the variance.
The constant “a” is found by taking mean values in equation (1).
y =a+bx
or a=y-—bx (2)
The ionic relationships obtained in this manner are:
Cl = 058H +914 (r= 0-77) (3)
Na‘ = —0:521H + 86:9 (rx =-0-83) (4)
K* =—0-083H + 26-46 (1 = —0-27) (5)
The nomenclature used to express the Hollander two-component theory is
shown in Table 4.
€. W. MILTON, D. J HINGSON AND E. P GEORGE 65
If the two compounds are isotonic
ptq=b+e+d (6)
The relations between the ionic concentrations have been outlined by James
(1957), and if x is the proportion of the parietal cell component in gastric
juice, the relevant equations are:
| Cl = bx + (c+) (7)
Na* = (c+b) (1-x) (8)
K* = qx+d (1-x) (9)
H’ = (pt+b) x-b (10)
We require to know the concentrations of Cl, Na and K’ in terms of H’ as
in equations (3), (4) and (5). Therefore by eliminating x from equations
(7), (8), (9) and 10 we obtain:
b b’
Cl =—— H+ ap (Gar
ptb ptb (11)
(c+b) (c+b)
Na* = -H +
(ptb) (pt+b) (12)
(d+q) bq+pd
K =-H op one
(p+b) p+b (13)
c+d c+d
H* = K ——_-+-q
d=qis id= (14)
The volumes of Cl, Na‘, K* and H’ represent the total concentration of each
ion in whole gastric juice. By solving the equations relevant for each ion,
e.g., (11) and (3) for Cl, and (12) and (4) for Na’ a series of values is
obtained as in Table 5 which are in best agreement with the Hollander theory.
5 TABLE 3
Maximum concentration of Na, Cl, H and K at mEq/l. and of
Pepsin < 103 units/ml. +S.D. obtained from four isolated cardio-
gastric glands and four total gastric pouch experiments
Whole Stomach Cardiogastric Gland
(Group 1) (Group 2)
H 129-04 4-6 GoO2 OXNar
Cl 170:5+ 2-6 170-5+ 5-0
Na 16-34 3°8 24-74 10:5
30-0+ 4:-8 19-9+ 4-9
12 490-7+4137-0 624-7+4232-3
In these results the parietal and non-parietal components are not isotonic.
The ionic relations calculated from equations (11) to (14), using the
values in Table 5 are:
Cl = 058H +168 (15)
Na‘ =-0584H + 96 (16)
K* =-0-08385H+ 14 (17)
These relations are closest in the equations (8) to (5) in the present analysis.
For comparison, we also found the best fit on the assumption that the total
molarity of the non-parietal secretion was 170 mM., making it isotonic with
the parietal secretion.
E
66 STOMACH CAPACITY AND CARDIOGASTRIC GLAND OF THE WOMBAT
The best fit was found for the following values:
NaHCO; b = 120 mM.
NaCl c= 35
KCl d= 15
170
which give the following theoretical relations:
Cl = 0-42H +100 (18)
Na* =-0:54H + 90 (19)
K* =-0-052H+ 9 (20)
which again agree adequately with the experimental results (compare
equations 3-5 with 18-20).
TABLE 4
Parietal cell secretion Non-parietal cell secretion
(Concentration) (Concentration)
HCl Pp NaHCO, b
KCl q NaCl ©
KCl d
The figure of 170 mM. concentration assumed for both parietal and non-
parietal solutions in the last analysis is not inconsistent with the measured
osmolarity of 281-4 ml./l. (Table 1), since the latter figure is weighted strongly
by the fully ionised solutes by a factor of two. However, the non-parietal
figures given in Table 5 are probably inconsistent with the measured
osmolarity.
TABLE 5
Parietal cell secretion Non-parietal secretion
(Concentration mEq/1.) (Concentration mEq/1.)
HCl p 167 NaHCO, b 232
KCl gq 0 NaCl c 0
KCl d 33-4
Total concentration 167 265-4
The results were analysed to determine whether it was justified to group
together the results for whole stomachs and for glands. From this analysis it
appears that the Cl/H’ and the Na‘*/H’* relations are the same for both
gland and for stomach, but that the K’/H’ relations are different. We find
for the cardiogastric gland alone
K =-0:30H +54 (r=-0-52)
whereas for the whole stomach alone
K =-0-14H + 37-2 (x =-0-47).
The separation into two sets is seen to improve the correlation. These results
taken together with the relations (3) and (4) give the Hollander analysis of
the secretions (Table 6), which suggests that the parietal cells of the stomach
secrete a small amount of HCl (20 mM.), whereas the parietal cells of the
cardiogastric gland do not.
DISCUSSION
The wombat is a member of the Phascolomydiae family of the marsupials.
The primitive forms of the marsupial separated from the ancestors of the
present day mammals, in the remote past, probably about 70 million years ago.
“G. W. MILTON, D. J HINGSON AND E. P GEORGE 67
Although in the intervening periods the marsupial has undergone considerable
changes, especially in the larger Australian varieties, there is still an enormous
eulf between the marsupial and the placental mammal. It is therefore of some
interest to compare the secretory ability of this animal with that of more
usual laboratory animals and man. On the whole the function of this “antique”
stomach shows a remarkable resemblance to that of man. The stomach
responded to histamine stimulation with a brisk ouptput of gastric juice and
the concentration was only slightly lower than the maximum concentration
achieved by man, dog and cat. The relationships of the different ions to one
another were similar to those of the mammals. The Hollander two component
theory still broadly fits the secretory pattern of this animal. The fit between
theory and the observed results was not so close as has been observed in the
cat (Milton et al., 1963). This may be partly explained by the fewer number
of experiments and the more restricted scatter of the observed ionic concentra-
tions. The electrophoretic pattern of the gastric juice also resembled that of
man.
TABLE 6
Parietal cell secretion Non-parietal cell secretion
Gland Stomach Gland Stomach
HICH 5% 167 167 INGECOR ae 100 100
KCI Rt 0 20 NaCl Be 35 35
KCI ss 55 55
The close correlation between K* and pepsin, if confirmed in man, could
be a useful indication of peptic concentration in human gastric function tests,
as it is easier to measure K* concentration than pepsin.
Much has.been written about the “maximal” gastric secretory capacity.
These findings show that in this animal, when the stomach is secreting at
close, to the limit of its power as a response to histamine, then a considerable
boost in secretion can be achieved by the addition of insulin to the stimulus.
Stimulation with combined agents may therefore give a higher maximal level
than the use of one agent alone.
The finding that the cardiogastric gland secretes juice of about the same
eoncentration as the body of the stomach agrees with the histological findings
that the cells composing the cardiogastric gland are similar to those of the
corpus (Hingson and Milton, 1968). It was not feasible in these experiments
to compare in detail the output of the cardiogastric gland to the rest of the
stomach in terms of ml. per minute; a large number of animals would be
required to do this and it would be necessary to standardise the size of the
stomach and gland in each case. But in the intact animal the cardiogastric
gland must secrete a large proportion of the gastric juice. The reason for the
slight difference in the relationship between K°/H™ in the whole stomach and
the gland is not clear, but it could be related to the very large number of
chief cells in the gland area and their intimate relationship to the parietal
cells, so that during acid secretion in the gland the cells extract some K*
from the adjacent chief cells.
Acknowledgements
We are grateful to Professor John Loewenthal for his help and encourage-
ment during this work. Part of the expenses for one of us (D.J.H.) was
defrayed by a grant from the M.C.P. Pure Drug Company, Wembley,
Middlesex. We are particularly grateful to Dr. M. Griffiths for help with the
preparation of this paper.
68 STOMACH CAPACITY AND CARDIOGASTRIC GLAND OF THE WOMBAT
References
Harrison, D. D., 1964.—Plasma pepsinogen as a test of gastric function. Proc. Aust. Ass.
clin. Biochem., 1(3): 75.
Hincson, D. J., and Minton, G. W., 1968.—The mucosa of the stomach of the wombat
(Vombatus hirsutus) with special reference to the cardiogastric gland. Proc. Linn.
| Soc. N.S.W., 93: 69.
Ho.tiAnper, F., 1932.—Studies in gastric secretion. IV. Variations in the chlorine content
mot gastric juice and their significance. J. biol. Chem., 97: 585.
Home, E., 1808—An account of some peculiarities in the anatomical structure of the
wombat, with observations on the female organs of generation. Phil. Trans. roy.
Soc., 98: 304-312.
Houssay, B. A., 1955—‘‘Human Physiology.” McGrath, M., New York (2nd KEd.), p. 7.
James, A. H., 1957.—‘‘The Physiology of Gastric Digestion.” The Williams and Wilkins
Company, Baltimore, p. 50.
JoHNSTONE, J., 1898.—On the gastric glands of the marsupialia. J. Linnean Soc., 27: 1-14.
LEHMANN, J., 1939.—Elektrometrische Mikrobestimmung von Chlor in Vollblut, Serum
und Harn. Acta Paediat., 26: 258-267.
Mitton, G .W., 1962.—The behaviour of gastric epithelium under various circumstances.
Ann. roy. Coll. Surg. Engl., 30: 351-367.
Mitton, G. W., Sxyrine, A. P., and Grorer, E. P., 1963.—Histamine secretory tests in
cats. An evaluation of results based upon MHollander’s two-component theory.
Gastroenterology, 44(5): 642-653.
Muttier, E., 1942.—“Elektrometrische Masanalyse.” Kd. 6, Leipzig.
MacKkenzir, W. C., 1918.—“The Gastrointestinal Tract in Monotremes and Marsupials.”
Critchley Parker, Melbourne.
Nasser, E. S., 1953.—Gastric secretion in the beaver (Castor canadensis). J. Mam-
mology, 34: 204-209.
Pierr, D. W., 1966—Personal communication.
Pieper, D. W., Stret, M. C., and Buriper, J. E., 1963——The electrophoretic pattern of
normal human gastric juice and of the gastric juice of patients with gastric ulcer
and gastric cancer. Gut, a: 236-242.
Smiru, J. R., Freyraac, F. C., Hurzinea, H., and CiarK, L. P., 1911.—Preliminary studies
on the structure and function of the cardiogastric gland of the beaver. Colorado-
Wyoming Acad. Sci. J., 3: 58.
Spector, W. S., 1956—‘‘Handbook of Biological Data.’ Editor Spector, W. B., Saunders,
Philadelphia.
THE MUCOSA OF THE STOMACH OF THE WOMBAT (VOMBATUS
HIRSUTUS) WITH SPECIAL REFERENCE TO THE CARDIOGASTRIC
GLAND
Dickson J. Hincson* and G. W. MiLrony
(Communicated by Dr. Mervyn Griffiths )
(Plates I11-v)
[Read 27th March, 1968 |
Synopsis
The specialised cardiogastric gland region of wombat stomach which is characteristic
of koala and beaver stomachs as well, is located on the lesser curve near the oesophageal
opening. The cardiogastric gland in the wombat is distinctive because of its complex
group of mucosal sacculations which open into the stomach lumen via 25 or 30 large
erater-like ostia. The mucosa of this gland contains long, straight, closely packed,
unbranched gastric glands composed of the cell types found elsewhere in the stomach,
with chief cells concentrated at the base of the glands. Parietal cells are present in
great abundance. Typical surface and neck mucous and argyrophilic cells are also
present. The bizarre cardiogastric specialisation in the wombat is thus not cytologically
a separate organ from the stomach. However, it does contribute greatly to the total
secretory cell mass of the stomach.
INTRODUCTION
The stomach of the wombat (Vombatus), koala bear (Phascolarctos) and
beaver (Castor) are of unusual anatomical interest due to the presence of a
convoluted mucosal specialisation, or cardiogastric gland on the lesser curve,
near the oesophageal opening. The first reported observations on the stomach
of a wombat were made by Everard Home (1808); since then only brief
descriptions of gross anatomical features of the stomach have been published
(Oppel, 1896; Mackenzie, 1918; Milton, 1962). Some histological aspects of
the cardiogastric gland region were studied by Johnstone (1898), but the
techniques of that day did not permit micro-photography of the cytological
organisation of the mucosa. This study was undertaken in order to clarify
the organisation of this unusual stomach.
MATERIAL AND MrtrHops
Twenty-four adult wombats of both sexes (species: Vombatus hirsutus)
were trapped in rural areas of Victoria and New South Wales. The animals
were killed and the stomach was removed within five minutes of death,
distended with 10% formal saline and immersed in this fixative. One stomach
was sectioned and blocks placed in osmic acid-zinc iodide fixative, to stain
the postganglionic unmyelinated parasympathetic nerve fibres (Maillet, 1963).
Material for light microscopy was embedded in paraffin wax and sections 7-8
microns in thickness were stained by various methods. These included
haematoxylin and eosin, periodic-acid-Schiff reaction (PAS), block silver
impregnation (Masson, 1928), thionin and methylene blue. An incubation
time of 36 hours was found preferable for the block silver impregnation.
* Research Student, Johns Hopkins University, Baltimore, U.S.A. Present address:
Department of Anatomy, Harvard Medical School, Boston, Massachusetts, U.S.A.
+ Department of Surgery, University of Sydney.
This study was performed during an elective quarter from Johns Hopkins Medical
School. Partial financial support was received from Smith, Kline and French Labora-
tories, Sydney, and M.C.P. Pure Drugs Limited, Wembley, Middlesex.
PROCEEDINGS OF THE LINNEAN Society or New SoutH WALES, VoL. 93, Part 1
70 MUCOSA OF THE STOMACH OF THE WOMBAT
OBSERVATIONS
General Features of Wombat Stomach
The wombat stomach resembles that of the human in shape, and thus the
usual regional nomenclature will be used: cardia, fundus, corpus and pylorus.
The cardiogastric gland is found immediately distal to the cardiac region,
hence its name (suggested by Smith et al., 1911) for the analogous structure
of the beaver.
The stomach wall varies considerably in thickness, from 0-6 mm. in the
fundic region to 2-0 mm. in the corpus and pyloric region. The cardiogastric
gland is the notable exception, as the thickness of the stomach wall in this
region varies from 4 to 8 mm. The empty stomach measures 30 to 35 cm.
along the greatest length from fundus to pylorus. It is thrown into several
longitudinal folds which appear to converge on the cardiogastric gland except
at the pyloric border. These disappear when the stomach is distended.
The cardiogastric gland can be identified externally on the lesser curve
by several criteria:
(1) Its bulging position just distal to the oesophageal-gastric junction.
(2) A ruddy brown colouration due to the skeletal muscle fibres which
run over it from the oesophagus, parallel with the longitudinal axis
of the stomach. These fibres terminate, though not abruptly, near
the pyloric border of the gland.
(3) A fold of lesser omentum and large stomach vessels which are seen
crossing the stomach wall, just distal to the gland. However, the
eland borders are not sharply outlined on the serosal surface.
The cardiogastric gland, on the luminal side, is distinguished by 25-30
crater-like outlets into the lumen of the stomach (PI. m1, Figs 1 and 2).
‘These outlets, as shown in Figs 1 and 2 are convoluted within; they are not
simple ostia. The trapezoid shape of the gland is well defined by these outlets,
although convolutions of mucosa underlie the surface epithelium for a few
millimetres peripheral to the outlets. The gland has its axis on the lesser
curve; it converges to its narrowest dimension at the oesophageal opening.
The mean surface area of these “gland trapezoids” as measured in six wombats
was 15-0 em?. The outlets are often missing from the axis itself with only
simple ridging present instead. Occasionally an outlet can be seen somewhat
peripheral to and isolated from the main mass of the gland.
Regional Organisation of mucosa
The Cardia.—Typical cardiac, or mucus-secreting glands, are present in
a narrow ring-shaped area (2 mm.) surrounding the oesophageal opening.
Parietal cells begin to appear in the more peripheral part of this band. Only
mucous cells are present immediately adjacent to the oesophago-gastric
junction. The cardiogastric gland adjoins the cardiac area.
The Fundus.—The gastric glands in the fundus are much shorter and
more coiled than in other regions. The mucosal thickness here is only about
one-third that of the corpus and cardiogastric gland. Surface and neck mucous,
parietal, chief, and argentaffin cells are all present, but the chief cells are
much more abundant, relative to parietal cell population, than in the corpus
and cardiogastric gland. Chief cells almost exclusively occupy the basal half
of the individual gastric glands, with only occasional parietal cells or mucous
neck cells wedged in among them. The luminal half of the glands is composed
of surface mucous cells.
The Corpus.—This region is composed of long, straight, simple gastric
glands. They are densely packed, parallel to each other, and perpendicular
to the muscularis mucosae. The glands contain all five major cell types, and
DICKSON J. HINGSON AND G. W. MILTON ral
and empty into gastric “pits” or foveolae. The distribution of cells and
thickness of mucosa closely approximates the situation in the cardiogastric
gland, and will be described under that heading. The only difference between
this region and the cardiogastric gland is that here the mucosa forms a simple
lining sheet without convolutions and impocketings. The corpus mucosa blends
gradually into the mucosa. of other regions.
The Cardiogastric Gland.—The mucosa of the cardiogastric gland is
folded upon itself in an elaborate manner, so that sac-like impocketings
result. The nature of these impocketings is best understood by sections
through the edge of the cardiogastric gland, where the mucosa is beginning
to fold under itself (Pl. 11, Fig. 3). The sacs thus formed are simple,
branched, tubular, and do not anastomose. Muscularis mucosae closely invests
these sacs and their branches. The ostia of the sacs are 1-5 mm. across, and
represent the 15-30 crater-like openings seen macroscopically. Because of the
large volume occupied by the impocketings relative to the smaller area required
for their ostia, the underlying mass of sacs bulges beyond the outer rows of
ostia. Thus it is that the peripheral rim of the cardiogastric gland underlies
a simple epithelial sheet that does not bear openings (PI. 11, Fig. 3). As
Fig. 3 also shows, the submucosa sweeps up among the sacs, bringing in its
areolar tissue nerve, lymph and blood supply to the sacs, as well as some
smooth muscle slips which are probably derived from the muscularis mucosae
and externa. The muscularis and serosal coats simply pass underneath the
whole mucosal mass, without investing individual sacs. The collection of longi-
tudinally oriented striated muscle fibres, just deep to the serosa, is easily
seen histologically (Pl. 111, Fig. 4). For a more detailed description of the
muscularis layers the reader is referred to Johnstone’s account (1898).
A section cut transversely across the lesser curve (the axis of the gland)
“mImay show a bifid ridged median leaf of submucosa, with no underlying sacs,
and correspondingly thickened submucosa and muscularis layers (Pl. 111,
Fig. 4).
The mucosa of the gland is similar to that of the corpus in cytological
organisation. While its single-sheet thickness (0-63 mm.) is approximately
that of the corpus, the mucosa is piled up so that the mean total thickness of
mucosa in a given section is four to ten times that of any other stomach region.
The individual glands are simple, straight and densely packed, opening
near the lumen of the stomach into foveolae (Pl. tv). These contain the
five principal cell types found in other mammalian stomachs. The chief
cells are concentrated in the basal fifth of the glands, with occasional parietal
cells wedged among them. A few argentaffin cells are near the base of some
of the glands. A large number of parietal cells are present in every gland
in its middle three fifths with mucous neck cells or, occasionally, surface
mucous cells interspersed. Approximately 500 cells constitute a single gland.
Connective tissue, capillaries, fibroblasts, and a few smooth muscle fibres
fill in the spaces between glands. The capillary piexus is particularly rich
in the muscularis mucosae and near the gastric foveolae, just beneath the
surface mucous cells. Large, thin-walled vessels, packed with blood cells, are
often seen between foveolae.
The Pylorus.—The mucosa of the corpus gradually loses its parietal and
chief cells as the pyloric opening is approached, although argentaffin cells
remain. The epithelium becomes thinner, and foveolae eventually extend half-
way to the muscularis mucosae. The glands become entirely composed of
mucus-secreting and argentaffin cells. There is an intermediate zone, approxi-
mately 2 cm. wide, containing both corpus glands and pyloric glands. This
zone begins at the pyloric edge of the cardiogastric gland and continues
(2 MUCOSA OF THE STOMACH OF THE WOMBAT
around towards the greater curve. Near the pylorus the glands become widely
spaced, and considerable connective tissue, fibroblasts, and smooth muscle
fills the spaces between glands. The glands themselves are coiled and may
show simple branching. A suggestion of villi is seen in the few millimetres
immediately surrounding the pyloro-duodenal junction, with several gland
foveolae opening into each villus.
Cytology of the Gastric Glands—Except where stated otherwise, these
observations apply equally to cardiogastric gland and other regions of the
stomach. Five distinct mucosal cell types are evident; surface mucous, neck
mucous, parietal, chief and argyrophilic cells.
Surface Mucous Cells—Surface mucous cells line the foveolae of the
gastric glands and bridge the mucosa between the foveolae. They are tall,
columnar cells, with a height of 15 to 20 microns and a width of about four
microns. The nucleus is spherical and basally or centrally placed. A large
nucleolus may be present. These cells are easily identified in PAS sections
(Pl. v, Fig. 1) by the heavily striated mucous granules in the cytoplasm.
These granules are densely packed and are present in the apical half of the
cell. They do not encroach on the nucleus so as to flatten it, and usually a
clear zone is seen between nucleus and mucus accumulations.
Neck Mucous Cells—Neck mucous cells are found among the parietal
cells along the elongated neck of the glands. They are low columnar or
cuboidal and about 5 in diameter. The shape of these cells is variable, and
the cell may have a broad base and narrow apex, or vice versa. The nucleus
is basally located and flattened by an encroachment of mucous granules on
its apical side. Sometimes PAS-positive material is found on the basal side
of the nucleus.
Chief Cells ——Chief celHs are localised in the basal regions of the gastric
glands, and stain readily with basic dyes. They are cuboidal or low columnar,
and measures 5-10» in diameter. The nucleus is spherical and centrally
located. Accumulations of chromophil substance are present in the basal part
of the cell. This substance is distributed in parallel, concentric or radially
oriented rows, which give the cytoplasm a clearly striated appearance (PI. v,
Fig. 2). “Nebenkerns” (concentric spirallying ergastoplasmic rings) are
clearly evident in some cells. The striations often seem centered about or
actually attached to the nuclear membrane. The thickness of each striation is
0-2 microns or less. Accurate measurement is not possible because of the
limited resolving power of the light microscope.
The apical portion of the cell has a pale, bubbly or frothy appearance
suggestive of the zymogen granules characteristic of these celis. The vacuola-
tions are spherical and are about 1p in diameter. The top of the cell is
frequently seen bulging into the lumen of its gland tubule. This distension
is evidently caused by local accumulation of the secretory product. In certain
regions of the mucosa most chief cells are seen to be almost full of the
zymogen granules, with the ergastoplasm confined to the extreme base of the
cell. In other mucosal regions the reverse is true, with ergastoplasm occupying
the entire cell.
Parietal Cell.—Parietal cells are easily distinguished by their large size
(diameter 10—25y), and acdiophilic cytoplasm. They are confined mainly to
the middle three fifths of the gastric glands, but occasionally wedge among
both surface mucous cells and the basally located chief cells. Parietal cells
appear to have a slightly larger mean diameter in the cardiogastric gland
than in other stomach regions, ‘although this has not been aevaia alli verified.
Parietal cells are spherical or pyramidal in shape. The nucleus is large,
DICKSON J. HINGSON AND G. W. MILTON 73
centrally located and may contain a nucleolus. The cytoplasm has a definite
granular appearance. An intracellular canaliculus is present, staining nega-
tively in haematoxylin-eosin preparations. The canaliculus is sharply outlined
occasionally in PAS-treated material, as PAS-positive substances in the
canaliculus cause a definite purplish ring (Pl. v, Fig. 3). This canaliculus
surrounds the nucleus in a nearly completed “horseshoe”, the open ends of
which join the lumen of the gastric gland. The canaliculus follows a course
midway between nuclear membrane and external limiting membrane.
The Argyrophilic Cell—Argentaffin and other argyrophilic cells are
present throughout the stomach mucosa, though not in great numbers. Both
are demonstrated by the Masson stain. When present, they are usually basally
located in the gastric glands. Their size is difficult to measure, owing to the
great variety of shapes they assume (Pl. v, Fig. 4). Some cells are spherical
and are about 10u in diameter, others are greatly elongated (up to 30» in
length), with a narrow width (5) corresponding to the diameter of the
nucleus. The nucleus itself is spherical and may be centrally placed in some
cells. Not infrequently the nucleus may be centered at one end of the cell,
with most of the cytoplasm concentrated around it. In such cases a long
tapering extension of cytoplasm extends in one direction away from the
nucleus, thus giving the cell a flask or club-shaped appearance. This extension
may narrow considerably until it resembles a thread-like neuron process,
0-24 in diameter. The argyrophilic cell seems to adapt its flexible shape to
directly conform with its neighbours, and does not necessarily abut on the
lumen of the gland tubules. The cytoplasm contains hundreds of small
granules which may be stained by silver impregnation methods. Many of these
granules are attached to the nuclear membrane.
Nervous Tisswe.—Anterior and posterior vagal trunks are present on the
oesophagus in its lower portion. These send branches to the cardiogastric
gland as well as to the remainder of the stomach. A definite “spray” of fibres
is often seen entering the cardiogastric gland.
Ganglia and nerve fibres were poorly or incompletely demonstrated in
haematoxylin-eosin and the various special stains which it was hoped would
distinguish them. However, the existence of the myenteric plexus of Auerbach |
in the muscularis externa was clearly demonstrated, with both nerve fibres
and large multipolar neurons present. The submucous plexus of Meissner
was not clearly seen.
DISCUSSION
Some earlier observers, as well as the present authors, have seen that the
cardiogastric gland of the wombat is not histologically a separate organ from
the stomach (Oppel, 1896; Johnstone, 1898). Its mucosa is composed of the
same types of secretory cells which are found in other stomach regions and
in other mammalian stomachs. The similarities of thickness and cell distribu-
tion in corpus mucosa and cardiogastric gland mucosa do not suggest a
unique function attributable to the cardiogastric gland.
The parietal cells of the wombat cardiogastric gland appear larger than
those elsewhere in the stomach, an observation which had been previously
made in beaver stomach (Nasset, 1953). However, we cannot verify in the
wombat any increased height of mucosa or density of parietal cells in the
cardiogastric mucosal lining as compared to neighbouring mucosa in the
stomach body; such an increase has been reported in the beaver (Nasset, 1953).
No comparisons based on those parameters have been reported for the koala
stomach. It must be recognised that the total amount of enzyme and acid-
secreting mucosa in the gland represents a very considerable fraction of the
74 MUCOSA OF THE STOMACH OF THE WOMBAT
total stomach mucosa. While the localisation of this mucosa has been carried
to an extreme in the wombat, koala and beaver, regional concentrations of
parietal cells have been observed in other mammals. In man, dog, cat and rat
the greatest density of parietal cell population exists in the stomach body
on the greater curve (Oi et al., 1958). The rabbit has the greatest concentra-
tion in the fundus, while the guinea pig has its greatest parietal cell
concentration in the same site on the lesser curve as does the wombat (Oi ez al.,
1958). The guinea pig does not possess a cardiogastric gland. Of interest is
the fact that in human embryos gastric pits and differentiating gastric epi-
thelial cells are first seen along the lesser curve, particularly in the oral part
(Salenius, 1962).
The subcellular features of the mucosal cells in the cardiogastric glands
or in other regions of the stomach were not unusual. However, the clarity
and organised patterns of the ergastoplasmic striations in the chief cells
were of some interest and are believed to be a well-defined rough endoplasmic
reticulum. These striations have been described in the chief cells and
pancreatic acinar cells of other mammals (Hagenau, 1958; Dalton, 1951).
Johnstone (1898) and Oppel (1896) have speculated about the significance
of the cardiogastric gland in mammalian evolution and the genetic differences
which may underlie its development. Oppel believed that the glandular
apparatus of the wombat and koala blossomed into the cardiogastric gland
in order to facilitate assimilation of large amounts of food. Thus the giand
is viewed as an accessory structure which in practice augments the limited
circumference of the stomach and represents a _ beneficial evolutionary
development. It is not certain, however, in what way the digestion of large
amounts of food is particularly enhanced by this gland. We have obtained
considerable quantitative data on the acid, enzyme, and electrolyte composi-
tion of sections from the cardiogastric gland. Further discussion of the
physiologic role of this anatomical specialisation is put forward by Milton,
Hingson and George (1967).
Acknowledgements
We are very grateful to Professor John Loewenthal, Professor of Sursery,
University of Sydney, for his encouragement and helpful criticism during this
work. We are also indebted to Mr. AS S. Cunningham and Mr. A. K. Carkeek,
of the Departments of Agriculture and Lands, respectively, of the State of
Victoria, for supplying the animals or stomachs thereof which were used in
this study. Professor K. W. Cleland and Dr. E. W. van Lennep of the
Department of Histology and Embryology, University of Sydney, offered
helpful advice and made available resources of their Department and technical
staff. Dr. J. I. Johnson of the Department of Physiology, University of
Sydney, also gave assistance in certain staining procedures. To all these we
are grateful. We are particularly grateful to Dr. M. Griffiths for his help
with this paper.
References
Daron, A. J., 1951.—Electron microscopy of epithelial cells of gastrointestinal tract and
pancreas. Am. J. Anat., 89: 109-133.
Hacenavu, F., 1958.—The ergastoplasm: its history, ultra-structure and biochemistry.
Internat. Rev. Cytol., 7: 425-438. :
Homer, E., 1808.—An account of some peculiarities in the anatomical structure of the
wombat, with observations on the female organs of generation. Phil. Trans. roy.
Soc., 98: 304-312.
JOHN STONE, J., 1898.—On the gastric glands of the marsupialia, J. Linnean Soc., 27: 1-14.
Maituet, M., 1963.—Le reactif au tetroxyde d’osmium iodure du zinc. JZ. Mikr. Anat.
Forsch., 70: 397-425.
Proc. LINN. Sag, N.S.W., Vol. 98, Part 1 PLATE III
£
The mucosa of the stomach of the wombat.
4
2
i
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e =
rh a“ oh ot
a me me
Appar hays Ags
i
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ate Se a
Proc. Linn. Soc. N.S.W., Vol. 93. Part 1
PLATE Iv
. Nee ote ro A Ss Pe §
eo fe Gs “ &,
S &
< Ri ‘or oye
ia!” Pie :
IG — aN
Ni, ee ov = P
The mucosa of the stomach of the wombat.
7
eanek
oe
se
‘—
Proc. Linn. Soc. N.S.W., Vol. 98, Part 1 PLATE v
The mucosa of the stomach of the wombat.
DICKSON J. HINGSON AND G. W. MILTON 75
Masson, P., 1928.—Carcinoids (argentaffin-cell tumors) and nerve hyperplasia of
appendicular mucosa. Am. J. Path., 4: 181-212.
Mixtton, G. W., 1962.—The behaviour of gastric epithelium under various circumstances.
Ann. roy. Coll. Surg. Engl., 30: 351-367.
Minton, G. W., Hineson, D. J., and Grorcr, E. P., 1968.—The secretory capacity of the
stomach of the wombat (Vombatus hirsutus) ata the cardiogastric gland. Proc.
Linn. Soc. N.S.W., 93: ???.
MACKENZIn, W. C., 1918. —“The. Gastrointestinal Tract in Monotremes and Marsupials.”
Critchley Parker, Melbourne.
NAsseET, E. S., 1953.—Gastric secretion in the beaver (Castor canadensis). J. Mammology,
34: 204-209.
Or, M., Sueimura, T., MoroyAma, A., KAwAmuraA, M., Komatsu, S., and Tortumi, T.,
1958 —Distribution of parietal cells of stomach in animals: dog, cat, rabbit, guinea
pig and rat. Jikei med. J., 5: 67-75.
OppPEL, A., 1896.—‘‘Lehrbuch der vergleichenden mikroskopischen Anatomie der Wirbel-
tiere. Teil 1. Der Magen.” Gustav Fischer, Jena, p. 291 and 402-403.
Satenius, P., 1962—On the ontogenesis of the human gastric epithelial cells. Acta
Anat., 50 (Supp. 1): 1-76.
SmitH, J. R., Freyrac, P. C., Hurzines, H., and CrarKk, L. F., 1911.—Preliminary studies
on the structure and function of the cardiogastric gland of the beaver. Colorado-
Wyoming Acad. Sci. J., 3: 58.
EXPLANATION OF PLATES III-V
PLATE III
Fig. 1. The stomach of the wombat turned inside out and distended, from EH. Home
(1808). The gland area is well shown on the lesser curve close to the oesophagus.
Fig. 2. Close-up photograph of the cardiogastric gland. The oesophagus is the
opening in the upper part of the picture. The edge of the antrum is at the extreme
bottom. The cardiogastric gland ostia spread out close to the oesophagus on the lesser
curve.
Fig. 3. Low power section, stained with haematoxylin-eosin through anterior edge of
wombat cardiogastric gland. A typical sacculation with its ostium is visible, resulting
from an impocketing of the stomach mucosa. Muscularis mucosae and submucosa invest
the sacs, whereas muscularis externa passes deep to the whole of the gland mass. x9.
Fig. 4. Low power section, stained with haematoxylin-eosin, of cardiogastric gland.
The lesser curve has been transversely cut, and the axis of the gland lies approximately
through the V-shaped median ridge in the centre of the photograph. Mucosal impocket-
ings and sacs are on both sides of this ridge, which is characterised by a thinner
mucosa than seen elsewhere .in the gland. Bundles of skeletal muscle fibres, transversely
cut, are just deep to the serosa. x 10.
PLATE IV
Section of cardiogastric gland mucosa, stained with PAS and counterstained
with haematoxylin. Gastric glands are seen to be simple and unbranched. Chief cells
are concentrated at the base of the glands and stain with haematoxylin. Parietal cells
remain unstained, while neck and surface mucous granules are stained intensely with
the PAS reaction. x 240.
PLATE V
Fig. 1. Transverse section through a gastric tubule from wombat cardiogastric gland.
The section has been stained with PAS and counterstained with haematoxylin. Tall
columnar mucous cells line the foveola, with dark-staining mucin granules evident in the
apical portion of the cells. An unstained parietal cell is seen wedged among these cells
at the upper left, communicating with the lumen through an intercellular cleft. x 2000.
Fig. 2. Section through the base of a gastric tubule in the cardiogastric gland,
stained with haematoxylin-eosin. Prominently striated ergastoplasm is seen in all of
the chief cells. The strands are oriented in a parallel, concentric, or radial pattern.
The cell in the centre of the microphotograph contains haemophilic strands which are
centered about or attached to the nuclear membrane. x 2000.
Fig. 3. Section through cardiogastric gland mucosa, stained with PAS and counter-
stained with haemotoxylin. The intracellular canaliculus of the parietal cell is
prominently stained. x 2000.
Fig. 4. A photomicrograph of an argentaffin cell from the cardiogastric gland. The
granules have been stained with Masson silver impregnation. Intensely silvered granules
fill much of the cytoplasm of the argentaffin cells, with other gastric epithelial cell types
almost entirely unstained. x 2000.
A REVIEW OF THE GENUS HALOCYNTHIA VERRILL, 1879
Patricia Korr
Zoology Department, University of Queensland
{Read 27th March, 1968]
Synopsis
The genus Halocynthia Verrill is reduced by synonymy to six closely related
species distinguished only by the condition of the gonads and the branchial and atrial
spines. Considerable variation in external appearance is demonstrated within a single
species.
Species occur in the littoral fringe of land masses and generally have a wide
latitudinal range. Their distribution appears to be limited mainly by deep waters.
The genus appears to be an ancient one and it may represent a relict of the Tethys
Sea fauna.
INTRODUCTION
The genus Halocynthia Verrill, is an homogeneous one, comprising only
a limited number of closely related species. These generally have a wide
cosmopolitan distribution from north to south along the sub-littoral fringe
of land masses in depths up to about 200 m. Geographic isolation is not
always a major factor in speciation and phylogeny of the genus is discussed.
An attempt is made in the present work to define the limits of intraspecific
variation and to clarify the taxonomy of the genus.
Genus: Hatocynruia Verrill, 1879
Type Species.—Ascidia papillosa Linnaeus, 1767.
Test produced into spines. Longitudinal glandular plications and
arborescent liver lobes present in the pyloric region. Gonads form, on each
side of the body, a parallel series of tubular ovaries surrounded by testis lobes,
each gonad terminating in a ? and ¢ duct directed toward the atrial opening.
On the left the gonads are in the gut loop and the ducts extend across the
descending limb of the intestine. A double series of languets present along the
dorsal line.
Species of this genus are present in sand, gravel or on rocks but never
in mud. The genus has been recorded, on several occasions, with the
phanerogam, Posidonia, which also flourishes on a sandy substrate (A.
papillosa in the Mediterranean and H. hispida in South Australia). Further,
the genus is invariably taken in fairly shallow water (up to 200 m. but
generally less) in sheltered waterways and estuaries where terrestrial run-off
and/or melting ice might be expected to affect the salinity of the water, e.g.,
Upper St. Vincent’s Gulf, Port Jackson, D’Entrecasteaux Channel, Akkeshi
Bay, Puget Sound, Massachusetts Bay, Gulf of St. Lawrence, Iceland,
Greenland, ete.
Key to the Species of Halocynthia
1 spines around apertures with’ secondary (spines 9.0.2.4. 00 oo ae eee 2
Spines around apertures without secondary spines .................. Pas ar 4
2(1) 1 or 2 gonads per side, not parallel, no spinules on shaft of spines..........
a pisten sere, sYellaiorist oye bepa eis IsEOS ee H. igaguri Tokioka
(Inland Sea, Japan)
Spinulesmon shaft sor spines: My tte een eee SND RE EY RENEE STS G80 0 3
PROCEEDINGS OF THE LINNEAN Society oF New SoutH WALES, VoL. 93, Part 1
I PATRICIA KOTT 0
3(2) More than 2 parallel gonads per side ................. H. hispida (Herdman)
(Hokkaido, Ceylon, East Australia and South Australia)
2 parallel gonads joining ventrally to formaU ........... H. spinosa Sluiter
(South Africa, Hast Africa, Red Sea)
4(1) 2 parallel gonads joining ventrally toformaU....... H. papillosa (Linnaeus)
(Mediterranean, Adriatic) ;
More than 2 parallel gonads joImeds wenitralllliven cps. * eealopench teie Len siereink ys ae 5
5(4) Surface of test raised into small spine bearing elevations (H. aurantium
CRallasy yi cs ape mar sttrans sperydionace tenons aleyracitaca APs caer eae sy coenccmen Peaeroa soho Coe nar, 6
Surface of test not raised into small spine bearing elevations H. roretzi (Drasche)
(Northern Japan, Japan Sea)
Gi@b) BRNO Thana citi corstean cu mou reste Peper eee eg we H. aurantium sub. sp. typicum
ING tPA ati Coe eis ceca sacra easiene csraloue apetmiarsl saniepae A. aurantium sub. sp. pyriformis
HALocyNTHIA HISPIDA (Herdman, 1881)
(Text-fig. 1)
Cynthia hispida Herdman, 1881, p. 61; 1882, p. 146; Cynthia crini-
tistellata Herdman, 1899, p. 34; 1906, p. 313; Halocynthia hispida; Kott,
1952, p. 288 var. typica; 1952, p. 284 var. crinitistellata; 1954, p. 129 var.
crinitistellata; Cynthia hilgendorfi Traustedt, 1885, p. 36; Oka, 1935, p. 436;
Halocynthia hilgendorfi; Hartmeyer, 1906, p. 6; Tokioka, 1959, p. 233; f.
rittert, 1962, p. 18; Halocynthia owstoni Oka, 1906, p. 42; Halocynthia rittert
Oka, 1906, p. 48; Halocynthia igaboja Oka, 1906, p. 45; Van Name, 1945,
p. 362; Halocynthia oka Ritter, 1907, p. 11; Ritter and Forsyth, 1917, p. 441;
Pyura okai; Hartmeyer, 1909-11, p. 134; Tethyum igaboja Huntsman, 1912,
pp. 114, 115, 136; Van Name, 1945, p. 362; Cynthia pachyderma Oka, 1926,
p. 559; Cynthia cactus Oka, 1932, p. 131; Halocynthia cactus; Tokioka, 1953,
p. 285; Rho, 1966, p. 213; 1966a, p. 366; ? Halocynthia simaensis Tokioka,
1949, p. 62.
Description.—The body is rounded, maximum diameter from 3 to 10 cm.
Individuals are often crowded together and the body becomes misshapen. The
colour is always red-orange. Posteriorly the test is produced into irregular
root-like processes. The test may be fairly thick and is always tough and
very leathery externally. The surface of the body is even; or produced into
tubercular prominences which are evenly distributed about 5 mm. apart over
the whole surface, or irregularly distributed. These tubercular prominences
are especially noticeable in the siphonal region where they are best developed.
Generally there seems to be a tendency to loss of the tubercular prominences
on the body with an increase in size although they do persist in the siphonal
region. There is also a tendency for the surface of the body to become
increasingly rough and wrinkled with increasing size.
Long spines of 2 to 3 mm. or more are often present either distributed
evenly over the body surface (C. cactus Oka, 1936); or supported by the
tubercular prominences singly or in groups of 2 to 3 (OC. crinitistellata
Herdman, 1899; Kott, 1952; C. hilgendorfi; Oka 1935; H. igaboja Oka, 1906;
Van Name, 1945). These long spines are absent altogether from some
Specimens (C. hispida Herdman, 1882; Kott, 1952. H. ritteri Oka, 1906;
Tokioka, 1962). In the present collection from St. Vincent’s Gulf all varieties
of test spine development are present:
Carickalinga Heads, 20 to 15 ft., “in caves and on vertical rock faces” :
(1) Even surface, globular body with slightly protruberant siphons. Only
few inconspicuous longer spinous processes from the test. Smaller test spines
covering the surface consist of 6 to 8 long radiating processes and a single
terminal process. Single specimen.
78 A REVIEW OF THE GENUS HALOCYNTHIA VERRILL
(2) Thickly distributed long spinous processes over the surface, longer
anteriorly, these are not always supported by tubercular prominences. Small
test spines covering the surface. Single specimen.
Off Port Stanvac, on steel wreckage:
Surface uneven, rounded tubercular prominences especially anteriorly,
Supporting single median, or several long spines. Smaller test spines with
‘6 to 8 stiff radiating processes. Single specimen.
St. Vincent’s Gulf, Posidonia beds:
Surface generally uneven with tubercular prominences, or irregular
transverse wrinkles. Long spinous processes present all over the body or
confined to the region around the siphons. Smaller test spines with 6 to 8
stiff radiating processes. Numerous specimens.
No constant condition in the distribution of the spines and tubercular
prominences has been observed in the specimens from any one area which
would suggest geographical subspecies: and specimens with spine arrange-
ment intermediate between those types described above are constantly
encountered. It is possible that the differences reflect to some extent environ-
mental factors (see below). Spines sometimes increase in length anteriorly
and in larger specimens become leathery. They have secondary spines
terminally and in 2 to 38 concentric rings along the shaft. The secondary
spines tend to lose their concentric arrangement as the spines become more
leathery and their distribution consequently becomes less regular. The shafts
of the spines are covered by regularly spaced minute spinules.
Over the whole surface of the test, between the longer spines, minute,
almost confluent papillae support 6 to 8 radiating spines or processes; or
they occasionally terminate in a single spine. The surface of these papillae
is also covered with spinules as on the shaft of the longer spines. These
papillae and the processes they support give to the surface of the test a
downy appearance. Always present around the apertures is a circle or
thicket of larger spines sometimes branched, and similar to those found else-
where on the test in the majority of specimens. They have secondary spines
terminally and in concentric circles along the shaft and they have spinules on
the shaft.
In the siphon linings scale-like swellings continuous with the small
Spine bearing papillae of the test extend in a single row down the folds which
correspond to the branchial and atrial lobes. These support one or more
small spines and in the furrows between the folds are reduced to single
conical spines or papillae.
Branchial tentacles vary from 8 to 16 with well-developed primary
branches supporting a fringe of minute secondary branches. Dorsal tubercle
forms a double spiral cone and rarely deviates from this condition. Dorsal
lamina consists of a row of pointed languets, closely placed; and to the
right of this a second row of similar languets but not so closely placed.
The branchial sac has 9 to 10 folds on each side of the body. The most
central fold on each side is often rudimentary. The maximum number of
vessels on a single fold is from 18 in a specimen of 1:8 cm. high up to 25 to 37
in specimens greater than 5 cm. There are from 1 to 3 longitudinal vessels
between the folds. Stigmata per mesh vary from 4 to 10 in individuals of
2 to 7 cm. H. cactus; Tokioka, 1953, 8 em. high had 20 to 23 stigmata in the
largest meshes but Oka’s specimens from the same locality and otherwise
identical with the former have a smaller number of stigmata in each mesh.
The gut loop is simple and closed. The ascending rectum extends
anteriorly to terminate in a smooth rimmed anus. Anterior glandular
3 PATRICIA KOTT 19
plications are present in the pyloric region and distal to these a collection
of arborescent liver lobules form a half ring around the gut usually on its
mesial surface. However, these may be displaced anteriorly around the
intestine.
Gonads vary from 2 to 10 parallel ovarian tubes on each side of the
body, directed toward the atrial opening. They are joined ventrally by an
antero-posteriorly oriented connective. Testes lobes are arranged along both
sides of the ovaries and in mature specimens extend as a continuous mat
Text-figure 1—Halocynthia hispida (from St. Vincent’s Gulf). 1 and 2. Specimens
from Carickalinga Heads; 38. Specimen from Posidonia beds; 4. Spines from thicket
around apertures; 5. Small spines on lobes of apertures; 6. Long test spine; 7. Small
test spines evenly distributed over surface; 8. Spines from inner lining of apertures;
9. Diagram of gut and gonads on the left.
between and on the parietal side of the ovaries. Testes ducts join on the
surface of each ovarian tube and form a vas deferens to open adjacent to
the oviduct. Occasionally gonads are missing or reduced on the right side
of the body. On the left they are present in the gut loop.
New Records.—From Posidonia beds, St. Vincent’s Gulf, South Australia,
5 fm.; 4 mls off Pt. Stanvac, South Australia, on steel wreckage, 15 fm.
80 A REVIEW OF THE GENUS HALOCYNTHIA VERRILL
Carickalinga Heads, 15 to 20 ft. in caves and on vertical rock faces, South
Australia. Coll. S. A. Shepherd.
Previous Records.—D’Entrecasteaux Channel, Tasmania, 5 fm. (Kott,
1952) ; off Maria Island, Tasmania, 174-155 m., 676-128 m. (Kott, 1954) ; Bass
Strait, 38-40 fm. (Herdman, 1882); Port Jackson (Herdman, 1899; Kott,
1952) ; Ceylon, 6-9 fm. (Herdman, 1906) ; Hokkaido, Japan (Traustedt, 1885;
Oka, 1906); Honshu, Kyushu, Japan (Oka, 1906, 1932, 1935; Tokioka, 1949,
1953, 1959, 1962) ; British Columbia to California, 10-90 fm. (Ritter, 1907;
Huntsman, 1912, 1921; Van Name, 1945).
Distribution.—In 5-90 fm. in the Pacific Ocean from Hokkaido to
southern Australia in the west and from British Columbia to California in
the east. The species has not been recorded from New Zealand; nor from any
other islands in the Pacific and its spread may be limited by deeper waters.
Records are lacking from the Malayan Peninsula, Indonesia, west and north-
east Australia. Further collecting may establish some continuity between
the Japanese, Ceylon and Australian specimens as there is no morpho-
logically stable characters which might suggest isolated communities in these
areas. Nor is its recorded distribution continuous across the north Pacific
Ocean from Hokkaido to British Columbia.
Habitat—Van Name (1945) describes the species as present on a sandy
or gravelly bottom. The present specimens are described as “thick in
Posidonia beds’; ‘vertical rock faces”. Specimens from d’Entrecasteaux
Channel were from scallop beds.
Remarks.—Externally therefore specimens of this species vary in external
appearance and may be characterised as belonging to the following types:
(a) Long spines absent from body of the individual; surface even:
Cynthia hispida, Herdman, 1882, Bass Strait; Halocynthia hispida var. typica
Kott, 1952, d’Entrecasteaux Channel; Halocynthia ritteri Oka, 1906; Tokioka,
1962, Japan.
(b) Long spines randomly distributed over the body of the individual;
surface even: Halocynthia cactus Oka, 1932; Tokioka, 1953, Sagami Bay,
Japan; Halocynthia igaboja Oka, 1906; Van Name, 1945, Japan and East
Pacific.
(c) Some spines present on parts of the body (intermediate between (@)
and (b)): Cynthia hilgendorfi Traustedt, 1885; f. ritteri Tokioka, 1959, Japan.
(d) Surface of body raised into tubercular prominences supporting
longer spines: Cynthia crinitistellata Herdman, 1899; Herdman, 1906, Port
Jackson, Ceylon; Halocynthia hispida var. cruutistellata Kott, 1952, Port
Jackson; Kott, 1954, off Maria Island, Tasmania; Halocynthia hilgendorfi;
Oka, 1935, Japan.
It has not been possible to divide specimens demonstrating these various
conditions of the test into geographical sub-species. However, it is possible
that the variations occur in response to some environmental factor.
The species resembles H. spinosa Sluiter in the form of the longer spines
on the test; however, in the latter species the smaller spines which cover the
test and cause its granular consistency are supported on small scale-like areas
rather than papillae, similar to the condition in H. aurantium but distinct
from the homologous structures in HA. hispida. Cynthia crinitistellata
Herdman, 1906 from Ceylon has papillae rather than scales supporting the
test spines and despite its location geographically is undoubtedly a synonym
of H. hispida.
~ PATRICIA KOU 81
H. simaensis Tokioka is listed as a doubtful synonym of this species.
Although externally the specimen resembles larger specimens of the present
species the glandular plications of the pyloric region are subdivided into
lobes; and spines are absent from the siphonal lining. So far only a single
specimen is known. Further collection may confirm these characters as
indicating specific distinctions rather than individual abnormality or the
effects of age. The absence of gonads on the right is not necessarily significant
as the number of gonads, especially on the right, varies considerably.
Herdman (1899) considers Cynthia dumosa Stimpson, 1855, a very likely
synonym of the present species. However, Stimpson’s specimen was not the
characteristic orange-red colour of H. hispida; and was taken from Port
Jackson on a muddy substrate which is unusual for H. hispida.
HALOCYNTHIA AURANTIUM (Pallas, 1787)
(Text-fig. 2, (10-12) )
Ascidia aurantium Pallas, 1787, p. 24. (For further synonymy see
accounts of subspecies below.)
Description—Halocynthia aurantium (Pallas, 1787) from the north
Pacific and H. pyriformis (Rathke, 1806) from the north Atlantic both have
papillary swellings all over the test supporting pointed spines singly or in
groups. The spines, however, do not radiate as in H. hispida but are shorter,
project forwards, and have « central spine which is longer than those which
surround it. The species lacks the longer branched spines of H. hispida
although a circle of enlarged spines is present around the apertures with
minute spinules or barbs along the shaft. Other characters resemble those
in H. hispida. The two subspecies are distinguished from one another only
by the numbers of gonads on each side of the body: 8 to 7 for sub. sp.
pyriformis and 8 to 4 for sub. sp. aurantium. This, as observed by Van Name
(1945) and Arnbick (1928) does not constitute a very convincing distinction
and relationships are better indicated by subspecific than by specific rank.
HALOCYNTHIA AURANTIUM (Pallas, 1787)
sub. sp. typica
(Text-fig. 2, (11-12),)
Ascidia aurantium Pallas, 1787, p. 240; Cynthia pyriformis; Traustedt,
1885, p. 34 (part) ; Cynthia superba Ritter, 1900, p. 590; Pratt, 1916, p. 667;
Cynthia deani Ritter, 1900, p. 590; Halocynthia superba; Oka, 1906, p. 41;
Hartmeyer, 1903, p. 200; Halocynthia deani; Hartmeyer, 1903, p. 200;
Tethyum aurantium; Huntsman, 1912, pp. 114, 115, 1386; 19120, p. 173;
Redikorzev, 1916, p. 169 (part); Halocynthia aurantium; (part) + forma
koreana Hartmeyer, 1903, pp. 195, 200; Halocynthia aurantium; Ritter, 1913,
p. 448; + H. superba Michaelsen, 1919, p. 11; Hartmeyer, 1921, pp. 30, 33;
Arnback 1928, p. 84; Pratt, 1935, p. 748; Van Name, 1945, p. 362; Tokioka,
195i —p. 17; 1967, p. 219: Monniot, 1965, p. 115.
Records.— Korea (Hartmeyer, 1903) ; Hokkaido (Traustedt, 1885; Oka,
1906; Tokioka, 1951, 1966) ; Kuril Is. (Pallas, 1787) ; Vladivostok, Okhotsok
Sea (Redikorzev, 1916) ; Bering Sea, Bering Straits, Puget Sound (Huntsman,
1912, 1912a; Ritter, 1900, 1918); Alaska, Pribilof Islands (Ritter, 1913).
Distribution.—A continuous distribution is recorded from the Bering Sea
to overlap limits of H. hispida off Hokkaido and Korea in the north-western
Pacific and off north-west America south to Puget Sound.
Habitat—tThe subspecies is taken in waters from 10 to 180 m. on sand,
sand with stones and shells, and sometimes on rocks (Arnbick, 1928).
F
82 A REVIEW OF THE GENUS HALOCYNTHIA VERRILL
HALOCYNTHIA AURANTIUM (Pallas, 1787)
sub. sp. pyriformis (Rathke, 1806)
(Text-fig. 10)
Ascidia pyriformis Rathke, 1806, p. 41; Sars, 1851; Cynthia papillosa;
(part) Traustedt, 1880, p. 407; Non Gunnerus, 1765; Cynthia pyriformis:
Stimpson, 1854, p. 20; 1860, p. 1; Packard, 1863, p. 412; Binney, 1870, p. 17;
Dall, 1870, p. 255; 1872, p. 157; Morse, 1871, p. 352; Kiaer, 1893, p. 67;
1896, p. 12; Metcalf, 1900, p. 510; Hartmeyer, 1901, p. 49; 1915, p. 313, Rhabdo-
cynthia pyriformis; Verrill, 1879, p. 27; Whiteaves, 1901, p. 268; Michaelsen,
1918, p. 11; Hartmeyer, 1920, p. 127; 1921, p. 30; 1923, p. 168; Arnback,
1928, p. 33; Pratt, 1935, p. 748; Van Name, 1945, p. 359; Millar, 1966, p. 99;
10.
——
1 mm
1:0 mm
140mm
Text-figure 2.—Halocynthia aurantium subsp. pyriformis (after Millar, 1966).
10. Small test spines. Halocynthia aurantium subsp. typica (after Tokioka, 1951).
11. Small test spine; 12. Spine from thicket around apertures. Halocynthia spinosa.
13. Spines from around apertures (after Michaelsen, 1918); 14. Small test spines (after
Millar, 1962). Halocynthia papillosa (after Michaelsen, 1918). 15. Spine from around
apertures.
Tethyum pyriforme; Hartmeyer, 1914, p. 1103; Berrill, 1935, p. 257; Tethyuwm
pyriforme americanum Huntsman, 1912, pp. 112, 148; Berrill, 1929, pp. 46, 48;
1935, p. 269; Pyura pyriformis; Procter, 1933, p. 284; Cynthia nordenskjoldi
Wagner, 1885, p. 156; Herdman, 1891, p. 577; Cynthia papillosa; Jacobson,
1892, p. 156; Halocynthia arantium; Hartmeyer, 1903, p. 195 (part) ; Bjerkan,
1908 (part) ; Michaelsen, 1918, p. 11; Harant, 1929, p. 66; Pyura aurantium;
Hartmeyer, 1909-11, p. 1831; Van Name, 1912, p. 5382; Tethyum aurantium;
Redikorzey, 1916, p. 169; Pyura pectinicola Michaelsen, 1908, p. 262;
neat 1909-1911, p. 1341; Tethyum microspinosum Van Name, 1921,
p. :
= PATRICIA KOTT 83
Records.—Massachusetts Bay, Gulf of St. Lawrence, Labrador (Van
Name, 1912, Huntsman, 1912); Ellesmeere Land, west coast of Greenland,
Iceland (Traustedt, 1880, Arnback, 1928); Faroe Is., Spitzbergen, Barents
Sea (Redikorzev, 1916, Arnbick, 1926); north-western Norway (Arnback,
1928, Millar, 1966); Bergen (Rathke, 1906, Hartmeyer, 1901, 1925) White
Sea; Murman coast (Redikorzev, 1916).
Distribution.—A continuous distribution across the north Atlantic with
its most southern extent at Bergen in the east and Massachusetts in the
west. In the north it extends from Ellesmeere Land the west coast of
Greenland, Iceland, the Faroe Is., Spitzbergen and the Barents Sea. It is
therefore present much further to the north than H. aurantiwm sub. sp.
typica of which the most northern limit is the Bering Straits. There are no
records from further east than the White Sea.
Habitat.—tThe species is taken from rock, sand, stones and shell in waters
of 0 to 114 m.
HALOCYNTHIA SPINOSA Sluiter, 1905
(Text-fig 2, (18, 14))
“An Ascidia quadridentata L.” Forskal, 1776, p. 9; Halocynthia spinosa
Sluiter, 1905, p. 16; Michaelsen, 1918, p. 7; Pyura spinosa; Hartmeyer, 1909,
p. 1841; Pyura (Halocynthia) spinosa; Hartmeyer, 1912, p. 181; Halocynthia
spinosa f. defectiva Millar, 1962, p. 201; Halocynthia arabica Monniot, 1965,
Dp. aoe f. defectiva; Monniot, 1965, p. 121; ? Halocynthia sp. Harant, 1929,
p. 67.
Description.—In this species the longer spines on the body and in a
thicket around the apertures are similar to those of H. hispida, with terminal
secondary spines and secondary spines more or less in concentric rings along
the shaft. Spinules are also present on the shaft of these spines. However,
the species is distinguished by the U-shaped gonads on each side of the body
similar to the gonads of H. papillosa; and by the distribution and form of
the small test spines, supported on scale-like thickenings of the test 0-5 mm.
in diameter and consisting of a central spine 0-5 mm. long surrounded by
5 to 6 smaller spines 0-25 mm. long distributed around the border of the |
scale. These minute test spines are also similar to those of H. papillosa.
Records.—Red Sea, Gulf of Aden (Michaelsen, 1918); Somaliland
(Sluiter, 1905) ; Cape Province, South Africa, 0—13 m. (Millar, 1962); ? West
of Gibraltar, eastern Atlantic, 3745 m. (Harant, 1929).
Habitat.—The specimens from South Africa were all taken from rock.
There is no information on the type of substrate from which specimens from
other localities were taken.
Remarks.—Millar’s f. defectiva, taken from three different locations
between January and March, was so named due to the absence of gonads
on the right side of the body. While in Michaelsen’s specimens of the present
species gonads were absent on the right side. This absence of gonads from
one side occasionally occurs in other species of this genus but its significance
is not apparent.
The small specimen taken from 3745 m. west of Gibraltar (Harant, 1929)
has test spines typical of the present species, and, although the gonads are
not developed, probably represents an individual of this species. There are
no other records of H. spinosa from the Atlantic coast of Africa and this
single specimen may represent a relict population in deeper water.
The derivation of H. spinosa from the Mediterranean species H. papillosa
by the development of secondary spines on the longer spines of the test is
84 A REVIEW OF THE GENUS HALOCYNTHIA VERRILL
suggested by their otherwise close morphological similarity. Spread of the
ancestral species is unlikely through the Suez area as no continuity of the
marine environment existed there previous to the opening of the Suez canal
in the nineteenth century. Halocynthia papillosa has been known from the
Mediterranean since the seventeenth century and Forskal’s report of an
Ascidia quadridentata (synonym oi H. spinosa) is from the Red Sea in the
eighteenth century. The presence of H. spinosa in eastern Atlantic as indicated
by Harant’s (1929) specimen suggests a (not necessarily contemporary) circum
African distribution for the species. It is therefore most likely that the
ancestral species was continuous from the western Mediterranean and around
yia South Africa to the Red Sea. The Straits of Gibraltar subsequently
provided a sufficient barrier for the isolation of two distinct populations
representing the species H. spinosa and H. papillosa.
The significance of the close morphological relationship between this
species and H. hispida is also puzzling. In view of the wide distribution of
this and other species of the genus, there is no apparent isolating barrier
between H. hispida from Ceylon (Herdman, 1906) and the present species
from the Red Sea. The relationships of these and other species of the genus
are indicated in Text-fig. 3 and Table 1.
HALOCYNTHIA PAPILLOSA (Linnaeus, 1767)
(Text-fig. 2 (15))
Ascidia papillosa Linnaeus, 1767, p. 1087; Tethyum papillosum Gunnerus,
1765, p. 100; Cynthia papillosa; Savigny, 1816, p. 143; Heller, 1877, p. 249:
Lacaze Duthiers and Délage, 1892, p. 126; Roule, 1885, p. 180; Herdman,
1891, p. 576; Halocynthia papillosa; Hartmeyer, 1904, p. 322; Michaelsen,
1918, p. 10; Harant, 1929, p. 66; Harant and Vernieres, 1933, p. 24; Pérés,
1958, p. 161; Monniot, 1965, p. 113; Pyura papillosa; Hartmeyer, 1909, p.
1540; 1912, p. 181; Ascidia rustica Risso, 1826, p. 274; non Linné, 1772.
Description—tThis ‘species closely resembles H. aurantium in the
distribution and form of the small test spines, and in the distribution and
form of the larger spines which are present around the siphons. The larger
spines lack secondary spines but have spinules. However, the species are
distinguished by the U-shaped gonads of H. papillosa.
Records.—Western Mediterranean (Pérés, 1958, Harant and Vernieres,
1933); Adriatic (Heller, 1877); Atlantic coast of France (Harant and
Vernieres, 1933, Lacaze Duthier and Délage, 1892).
Habitat —From amongst coralline algae, Posidonia sp., sand and shell.
Remarks.—The morphological relationships between the present species
and H. aurantium in the north Atlantic and H. spinosa in the Red Sea are
indicated in Table 1.
HALOCYNTHIA 1GAGURI Tokioka, 1953
Halocynthia igaguri Tokioka, 1953, p. 20.
Description—This species is distinguished from all others by the gonads
which appear to be of a simple styelid type with tubular oviduct surrounded
by pyriform testes lobes. The long spines on the test have secondary spines
but no spinules and between these there are minute papillae evidently without
terminal spines as in H. hispida, H. aurantium and H. papillosa. There are
only 7 branchial folds per side and Tokioka has not described a double series
of languets along the dorsal line.
Record.—Inland Sea, Japan (Tokioka, 1953).
PATRICIA KOTT Sd
Remarks.—The condition of the gonads, number of branchial folds and
the dorsal lamina are not typical of this genus. However, there are the usual
glandular plications and liver lobes in the pyloric region.
HALOCYNTHIA RORETZII (Drasche, 1884)
(Styela?) Cynthia roretzii Drasche, 1884, p. 376. For further synonymy
see Tokioka, 1958, p. 282.
Description.—Specimens are up to 14 cm. long. This species has circles
of spines around the apertures similar to those of H. aurantium with spinules
but no secondary spines. The surface of the test is divided into scale-like
areas without spines, as in H. auwrantium. In older specimens large mam-
millary or finger-like processes develop from the test sometimes bearing a
terminal spine. There are very numerous branchial folds—up to 18 on each
side with 60 to 70 longitudinal vessels on each fold. Gonads are numerous—
( tomual::
Younger specimens have neither mammillary nor finger-like processes
and no polygonad scale-like areas; the surface has instead a number of
spines with spinules similar to those present around the apertures of the
adult.
TABLE 1
Morphological relationships between common species of the genus Halocynthia
Gonads
U-shaped Many parallel
gonads gonads
H. papillosa H. aurantium
Absent Mediterranean North Atlantic
: North Pacific
Secondary spines
on Test spines
H. spinosa H. hispida
Present South Africa North Pacific
Red Sea Indo-Malayan
For fuller description see Tokioka, 1953, p. 282.
Records —‘Coasts of Hokkaido, Honsyu, Sikoku and Kyusyu but not
from south coast of Sikoku and Kyusyu .. . also distributed throughout the
coast of Tyosen and the coast of Shantung Peninsular in North China.”
(Tokioka, 19538, p. 285.)
Distribution.—In fairly shallow water in a limited area around Japan
and the northern part of the Japan Sea.
Remarks—tThe species is probably related to H. auwrantium and its
distinguishing characters are largely a result of the greater development of
those characters than occurs in H. aurantium.
PHYLOGENY
(Table 1, Text-fig. 3)
The four most widespread species of this genus present interesting
morphological and geographical relations. From the condition in the
Mediterranean species H. papillosa with U-shaped gonads, simple spines, and
spine-bearing scales on the test, H. awrantium in the north Atlantic is
differentiated by development of the gonads; and H. spinosa from South
Africa and the Red Sea develops secondary branches on the test spines but
retains the U-shaped gonads. This distribution of closely related species
86 A REVIEW OF THE GENUS HALOCYNTHIA VERRILL
suggest that this genus represents a relict of Tethys Sea fauna. The extension
of H. aurantium into the north Pacific through the Bering Straits probably
occurred later. However, H. spinosa, radiating from west of the Medi-
terranean southwards around Africa and north into the Red Sea could have
been a Tethys component of the tropical Atlantic-West Pacific fauna. The
combination of characters found in H. hispida may result from H. aurantium
by the development of secondary spines, or from H. spinosa by the increase
in the gonads. The distribution of H. hispida in the north Pacific and Indo-
Malayan areas, as well as its morphology, is intermediate between 4H.
aurantium and H. spinosa and its origin is probably more recent than either
of the latter species.
; = Hhispida A
~ ispida
rs H. spinosa A
NS H.aurantium @
NG: H. papillosa tO)
H. rorefzii a
SOUTH AMERICA
NORTH AMERICA =:
NEW
ZEALAND
Text-figure 8. Map showing world distribution of species of Halocynthia.
Acknowledgements
The specimens of H. hispida from St. Vincent’s Gulf are part of a
collection made by Mr. S. A. Shepherd of the South Australian Museum.
The author is indebted both to Mr. Shepherd and to the Director of the
South Australian Museum for the opportunity of examining the collection.
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A NEW BDELLOURID-LIKE TRICLAD TURBELLARIAN
ECTOCONSORTIC ON MURRAY RIVER CHELONIA
LAURENCE R. RICHARDSON*
[Read 24th April, 1968]
Synopsis
A new genus is provided for a small acephalous, sedentary turbellarian with
posterior adhesive discs, a posterior male complex, and general form resembling
Bdelloura candida, a maricolous triclad on the gills of Limulus. It differs significantly
otherwise so that it is a novelty in the Maricola as well as the Paludicola, but is
provisionally placed in the latter. ‘“‘Perigrinatic”’ is proposed for consortism other than
parasitic, symbiotic, and commensal.
This paper describes a triclad turbellarian found in the limb-pits of
turtles from a lake near Griffith, N.S.W. As freshwater triclads these would
be expected to be Paludicola, but they possess in addition to an anterior
adhesive pad, longitudinal marginal adhesive bands and paired posterior
adhesive discs. They are acephalous and the single pair of eyes is in a
posterior position. These and other features are exceptional in the Paludicola,
seen in some Maricola but as yet no Maricola are known from freshwater.
The copulatory bursa is situated at the anterior end of the male terminal
reproductive organs, dorsolateral to them not fully anterior as in Paludicola,
and since the genital pore is posterior, marginal, located between the posterior
adhesive dises, the “probursal” condition is possibly secondary. I place these
animals provisionally and with much reservation in the Paludicola.
The relationship to the turtles does not fall into the usual consortic
categories when these are used in their proper sense, and I propose that it be
termed perigrinatic, in the sense of Virgil: “one travelling in a foreign land”.
There is nothing to indicate that this triclad may not be free-living. It is not
parasitic in the sense that it draws nourishment from the host, as does the
Sanguivorous leech, g. Placobdella, which is found with it. It is not symbiotic
in the correct usage of this term. There is nothing to indicate it benefits
the host or benefits from it. It does not share the food of the host since it
is microphagous, and so should not be termed commensal even recognising
that this category has become the catch-pot for any kind of consortism which
is neither parasitic or symbiotic.
Four specimens came to me from Dr. R. E. Barwick who found them
and others along with many egg-capsules in the limb-pits of Hmydura
macquartt and Chelodina longicollis taken at Lake Wyangan, near Griffith,
N.S.W. during the course of experimental fishing operations carried out by
officers of the N.S.W. Inland Fisheries Research Station towards the end of
May, 1967.
The stalked egg-capsules immediately recalled those of the bdellourid
Syncoelium and of some other Planariidae. The animals were fully bdellourid
like, even to the presence of two white bodies suitably placed to be the paired
copulatory bursae characteristic of this family. It was not until I had studied
serial sections that I could persuade myself that these animals could be
Paludicola, even though this also gave evidence of characteristics more suited
* 4 Bacon Street, Grafton, N.S.W., 2460.
PROCEEDINGS OF THE LINNEAN Society or New Sourn WALES, Vou. 93, Part 1
LAURENCE R. RICHARDSON 91
to the Maricola than to the Paludicola. Should it later be found that they
are maricolous, I see no genus there which will accept them (Hyman, 1951);
Grasse, 1961). The absence of a direct connection from the bursa to the
exterior excludes them from both the Bdellouridae and the Uteroporidae.
Otherwise there is the exceptional g. Puiteca in the Procerodidae which has
anterior and posterior adhesive discs and a posterior genital pore, but this
has the single pair of ovaries in the post-pharyngeal position and lacks a
bursa.
If paludicolous, the internal muscle layer of the pharynx includes a
circular layer uninterrupted by other muscle cells as is characteristic of the
Planariidae and Kenkidae. The latter is restricted to North America and
contains white, eyeless cave planarians with resemblance to the g. Phagocata.
There are only three genera (Hyman, 1951a), none suitable for the present
species. This presents a combination of characters which I have not found in
the various genera of the Paludicola (Hyman, 1951la, 6; Grasse, 1961), and
certainly not in the Planariidae. Accordingly, I propose a new genus as below.
MetTHODS
After preliminary study of the live specimens which provided information
mainly on the alimentary canal, the animals were narcotised in a drop of
water on a slide inverted over a drop of chloroform in a covered petri dish.
Then covered with a cover-slip, and 50% alcohol run in under the cover-slip,
the rate of flow being regulated to maintain just such pressure as held the
animal flat. A specimen was stained in acetic alum carmine, cleared in
glycerine, subsequently sectioned at 10 mu. and stained with Delafield
Haematoxylin and eosin.
BDELLASIMILIS, n. g.
Triclad bdellourid-like Turbellaria having a small anterior adhesive pad
continuous with narrow longitudinal marginal adhesive bands which expand
into two round ventral sucker-like posterior terminal adhesive discs;
acephalous; a single pair of eyes placed about 4rd of the length from the
anterior end; a single pair of ovaries close behind the eyes, prepharyngeal ;
copulatory bursa connects by a duct into the penis antrum; testes branching,
tubular, preocular and also lateral to the pharyngeal region; sperm vesicles,
paired, almost post-pharyngeal; sperm ducts join terminally into a common
median duct which connects to the penis where the cavity is central; no
bulbar cavity; genital aperture, posterior, median, marginal; no adenodactyl ;
no genito-intestinal connection; posterior limbs of intestine transversely
connected behind peripharyngeal chamber; egg-capsules, stalked, cylindroid.
Type Species—Bdellasimilis barwicki, n. sp. as follows:
BpDELLASIMILIS BARWICKI, Nn. Sp.
(hice)
A whitish or partly grayish, self-coloured, semitransparent, bdellourid-
like triclad of small size with a single pair of posteriorly placed eyes, and
with obvious paired posterior sucker-like adhesive discs.
Contracted (8-0 mm.) rather bluntly rounded anteriorly, the width about
2:0 mm. at the level of the eyes which are placed about 2-5 mm. from the
anterior end; maximum width of 3-0 mm. at the level of the posterior end of
the pharynx about 4:0 to 5-0 mm. from the anterior end. The margins then
curve obtusely so that the posterior end is obtusely rounded with the two
adhesive discs partly showing behind the margin of the body and separated
92 NEW BDELLOURID-LIKE TRICLAD TURBELLARIAN ECOTOCONSORTIC
by a wide and shallow notch where the elongated genital aperture opens
ventrally. The body is generally low convex above, flat below. When strongly
contracted, the surface shows numerous short striae which are transverse.
In extension it may nearly double its length, the preocular region becoming
elongate, tapering acutely to the narrow tip but the width and length of the
post-pharyngeal region are little changed from the contracted condition.
The anterior adhesive pad is transverse, short, only about 0:08 mm. in
length and continuous with the longitudinal marginal bands which are
0-10 mm. wide and run slightly medial to the lateral edge of the body which
overhangs them as though an eave. The bands expand posteriorly into the
adhesive discs which are 0°3 mm. wide and slightly longer than wide, with
narrow raised rims surrounding a flat central area which is richly supplied
with eosinophilic gland cells as also in the marginal bands and anterior pad.
Fig. 1. Bdellasimilis barwicki. A. Dorsal view of preserved specimen showing
distribution of chromatophores. B. Dorsal and C. lateral views showing the feeding
posture. Scale in mm.
The relationship of the rim to the disc is the same as the lateral edge of the
body to the marginal band. When the animal is detached from the substratum
and placed on its back, the rims of the discs may roll up, thicken, become
continuous, presenting the appearance of a single sucker which may be the
actual form of this organ: a single wide adhesive pad which appears as two
discs because of the emargination of the disc when attached to the substrate.
Seen with reflected light, the animal is whitish excepting for the black
and obvious eyes and pale brown chromatophores which are concentrated as
a longitudinal row on either side of the peripharyngeal chamber, the two
rows converging shortly behind the eyes and extending anteriorly between
them as a more diffuse median row nearly reaching the anterior end of the
body. Marginal bands of more widely scattered chromatophores run along
each side of the body. Between the paramedian row and the marginal row
there are scattered chromatophores Suggesting possible transverse bars and
~ LAURENCE R. RICHARDSON 93
others arranged over the posterior region of the body seem to be in radiating
lines. It is possible that the dorsum might be patterned when the chromato-
phores are extended. I have not seen this,
Alimentary System (Fig. 2D.)
-The ventral opening into the peripharyngeal chamber is about #ths of
the length of the body from the anterior end. It is transverse, narrowly
elliptical to slit-like, and the chamber wider and longer than the pharynx
and about 4th of the length of the body. The floor, walls and roof of the
chamber are much plicated internally and the whole chamber can be greatly
enlarged in the extended animal, especially dorsally as the roof is thin. The
pharynx, about {th of the length of the body is directed posteriorly and in
life, I did not see it protruded from the chamber. The pharynx is thick-
walled, the parenchyme towards the anterior end is rich in eosinophilous
gland cells which are densely packed also around the origin of the limbs of
the canal and along the anterior limb nearly to the level of the ovaries. There
are only one or two very short small lateral diverticula on the anterior limb
between the pharynx and the eyes. Anterior to the eyes there are some seven
or eight briefly ramifying elongate tubular diverticula on either side of this
limb, short in the sense that they reach only about half way to the margin,
and they diminish in length anteriorly. This limb reaches nearly to the
anterior end of the body.
The posterior limbs are nearly circular in section as they arch around
the anterior end of the peripharyngeal chamber, and depressed elliptical
lateral and posterior to the chamber. Lateral to the chamber, they carry a
few very small medial diverticula. There is a transverse canal joining the
two posterior limbs behind the chamber, followed by more numerous small
medial diverticula along the last portion which terminates bluntly near the
posterior margin at the level of the end of the penis. The lateral diverticula
are more lobed than tubular in their subdivisions, uniform in size along the
pharyngeal region and then diminish progressively in diameter and length.
Reproductive System (Fig. 2D, E). (Note: Dimensions given are taken from
sections. )
Testes, sperm ducts, vitellaria, and bursa could not be seen in the live
animal.
In sections, the testes and sperm ducts appear as a paired ramifying
tubular system anterior to the spermiducal vesicles and commencing about
0-75 mm. from the anterior end of the body so that the system is in part
preocular. The sperm ducts are somewhat coiling or tortuous, about 0-07 mm.
in width in the preocular region and slightly diminished in width as they
approach the spermiducal vesicles. The main ducts run just medial to the
longitudinal cords of the nervous system. If a side branch from the duct is
followed laterally, it closes off in simple blunt-ended tubules and there is
no spermatic tissue beyond this. From this, it seems the testes are ramifying
tubular lateral branches from the main duct, and openly continuous with
the duct. The histology of the contents of both tubules and ducts are the
same, both containing spermatocytes and spermatids. The ducts enter the
obliquely aligned somewhat ovoidal spermiducal vesicles located lateral to
the posterior end of the peripharyngeal chamber. These thin-walled vesicles
contain only mature sperm as also the following thin-walled narrow con-
voluted tubular sperm ducts which are about 0:03 mm. in diameter and
extend to the level of the penis bulb when each bends abruptly anteriorly
to join into a thick-walled transverse tube about 0:04 mm. in diameter and
94 NEW BDELLOURID-LIKE TRICLAD TURBELLARIAN ECOTOCONSORTIC
with a narrow lumen. From the middle of this tube, a thick-walled median
tube with a narrow lumen extends back and enters the penis. There is no
bulbar cavity. The canal is central in the penis which is about 0:04 mm. in
diameter and 0-12 mm. long, tapering, conical and protrudes into and almost
fills the proximal portion of the genital antrum. This antrum expands into
a second terminal chamber beyond the end of the penis and the slit-like
genital aperture opens ventrally from this second chamber. There is an
Fig. 2. Bdellasimilis barwicki. D. General morphology, composite based on the
living specimen, a stained whole mount, and serial sections. E. Free-hand reconstruction
from serial sections of terminal male organs, copulatory bursa and duct. F. Egg-capsule.
b.c., copulatory bursa; b.c.d., bursal duct; g.ap., genital pore; n.c., longitudinal nerve
cord; m.d., median duct; ov., ovary; pe., penis; pe.b., penis bulb; p.p.c., peripharyngeal
chamber; sp.d., sperm duct; sp.v., sperm vesicles; te., testis. Scale in mm.
4
4
LAURENCE R. RICHARDSON 95
elevation of the dorsal and lateral walls protruding into the antrum as
though dividing the proximal from the terminal chamber and leaving them
in communication only ventrally; but it is very short and the sections too
thick for me to determine the form of this structure.
The single pair of spherical ovaries are situated close behind the cerebral
ganglia and the eyes. They are 0:08 mm. in diameter and contain oocytes.
The vitellaria show in sections as irregular aggregations of cells chiefly in
the region of the ovaries and behind the ovaries, the greater part dorsal above
the anterior limb of the alimentary canal, and only some few cells between
the diverticula. There is no obvious pattern or distribution which might
indicate the path of a duct and I have been unable to detect a duct. The
oviducts could not be seen with sufficient regularity to indicate a path and
no terminal connections to the reproductive antrum or bursal sac or duct
could be found although the wall of both the proximal and terminal portions
of the antrum is thickly muscular. There are paired latero-dorsal brief
grooves on the inner face of the anterior end of the terminal portion of the
antrum suggestive of terminations of oviducts, but these grooves cannot be
traced into the muscular wall.
The copulatory bursa was not seen in either the live animal nor in
stained whole mounts. In sections it commenced as a slightly widened end
to a muscular cylinder about 0-1 mm. in length with a short bulbous caecum
on the dorsal aspect so that expanded it might become bilobate. It narrows
rapidly to a bursal duct of about the same length. This is thick walled and
with a narrow lumen. The duct extends briefly along the dorso-lateral aspect
of the penis bulb, enters obliquely to terminate by opening about half way
along and into the lateral aspect of the penis antrum. The thick walls of the
bursa and duct show no indication of the entry of oviducts and there are no
genito-intestinal connections.
The egg-capsules contain only a single embryo. There is an attachment
disc of the diameter or wider than the capsule, rather irregular in outline
and with some thin raised edges. The stalk is solid, short, cylindrical, its
length about equal to the diameter of the capsule which is 0-5 mm. wide and
about 1:3 mm. long, terminating obtusely with a minute spike. The colour
is brownish; the appearance, chitinoid; the wall, single; the surface, smooth.
A short cap detaches by a smooth-edged break running completely around
the capsule to release the young.
Type.—Whole mount stained specimen prepared by R. E. Barwick, Coll.
No. W 4174, Australian Museum, Sydney.
General Observations
Dr. Barwick noted that although he collected these turbellarians from
both Chelodina longicollis and Hmydura macquarti, there were few egg-
capsules and these only in the limb-pits of the latter in contrast to such
numbers in C. longicollis as to be described as dense masses in the order
of up to a hundred in one limb-pit alone, and also a few on the skin of the
legs elsewhere. On this evidence alone, it is clear there is a continued
association between B. barwicki and these Chelonia, for he found only a
few turbellarians which is suggestive of an extended period of capsule
deposition as is known in some triclads.
Of the four live specimens I received, two stayed persistently in the water
in the vial; two remained on the lower surface of the cap closing the vial.
They did not change position in 24 hours and seem to be strongly sedentary
in habit.
96 NEW BDELLOURID-LIKE TRICLAD TURBELLARIAN ECOTOCONSORTIC
The appearance is that of a glossiphonid leech when the body is extended
in contact with the surface, but it can be raised to the near vertical with
the animal erect on the posterior discs and then the lateral margins are
thrown into short rugae as though coarsely frilled. This attitude may be
‘sustained for five minutes and more. There is no movement such as the
respiratory movement in leeches when the body is raised clear of the surface.
When stimulated to move over a surface, the motion quite strongly
suggests euglenoid creeping. The prepharyngeal region is _ extended,
narrowed, but the pharyngeal and postpharyngeal regions are not greatly
reduced in width or length. The prepharyngeal region then widens and
shortens and the posterior region is drawn forward. It is not a rapid
movement. I did not see it exhibit typical smooth turbellarian progression.
Various attempts to persuade the animal to swim were unsuccessful. When
dropped into water, it sank slowly to the bottom, rolling up and unrolling
lengthwise and partly twisting the anterior portion of the body, but with
no control. In this it differs from Bdelloura candida which is a capable
swimmer (Verrill, 1892).
When detached from the surface and placed on its back, it is unable
to right itself. One was held in water in a watch-glass for three hours in
this position. During this time, it rolled up lengthwise repeatedly and
extended in attempts to find a hold for the anterior adhesive pad, but with
no success. It seemed to have no ability to twist the anterior portion of the
body sufficiently to obtain a hold. The posterior end of the body has even
less flexibility. The level of the water was reduced until the surface film
was within the reach of the animal but it made no attempt to utilise this
to re-orientate itself. In this inability to right itself, B. barwicki contrasts
with other Paludicola and with such Maricola as TI know.
It is a most difficult animal to manipulate. When detached from the
surface by a needle, the body is wrapped lengthwise or crosswise on the
needle adhering by the marginal bands, the anterior pad and posterior discs.
It cannot be shaken from the needle or displaced by even violent agitation
in water. As the animal is in fact firm bodied, it can be handled by using
two needles, transferring it from one to the other until it is moved to the
point of a needle, and it is unable to maintain a hold on this.
It is unusually light sensitive. An abrupt exposure to direct bright light
leads to strong contraction, and a gradual relaxation when the light is turned
off. When this is repeated several times, the animal acts as though condi-
tioned and remains contracted for ten to twenty minutes with the light turned
off. It does not respond to light from below even with very much higher
intensities than produces contraction with direct light above. The pigmented
optic cups open dorsally which apparently shields the light-sensitive retinal
structures from light from below. I have not found this response in Dugesia
or Curtisia spp. which I have known elsewhere.
B. barwicki has a distinct feeding posture (Fig. 1C). Attached by the
posterior adhesive discs, the body is raised and held parallel to the sub-
stratum. The postocular region is convex in profile; the extended preocular
region, concave above with the margins of the body rolled ventrally to form
a furrow which is open at the anterior tip of the body. The lateral margins
are undulate to irregularly rugose but without movement. The peripharyngeal
chamber is enlarged and obvious through the thin body wall roofing over it.
At intervals as frequent as ten minutes, the margins of the preocular region
are folded inwards, the body raised slightly as a whole, and the preocular
region curved ventrally and passed back under the ocular region and to the
LAURENCE R. RICHARDSON 97
back of the pharyngeal region, the whole as though forming a sac beneath
the peripharyngeal chamber. This attitude is held for a matter of a minute
or so, and then the former stance is resumed.
During this action, the opening of the peripharyngeal chamber is large,
twice and more the width of the pharynx, and the latter swings from side
to side, extends and shortens, as though probing all regions in the peri-
pharyngeal chamber. On one occasion, a considerable mass of unicellular
green algae was accumulated in the enlarged peripharyngeal chamber. The
margin of the enlarged opening into the chamber was thickened, as though
a band holding the mass of algae within the chamber while the pharynx
probed around and into it.
This manner of feeding suggested the possibility that there might be
something here of the nature of a ciliated mucous filter feeding mechanism
based on the mucus secreted from the anterior pad and marginal bands;
but I could not detect any signs of currents of water moving toward or away
from the animal such as would be expected in the presence of this type of
feeding mechanism. In fact, I was not able to detect any indication of
external ciliation in the live animal. The mass of uncellular algae in the
peripharyngeal chamber seemed to be a loosely formed accumulation and
the pharynx seemed to move freely into and through it with no indication
that the algae were trapped in mucus.
Acknowledgements
The specific name is given in appreciation of the assistance given to me
on this and other occasions by Dr. R. E. Barwick, Australian National
University. I desire to thank also Mr. N. Call of the same institution who
prepared the sections, and Dr. J. C. Yaldwyn, of the Australian Museum,
who has been most helpful on many occasions. I thank the Science and
Industry Endowment Fund for the loan of microscopic and other equipment.
References
Hyman, L. H., 1951a—North American triclad turbellaria. xii. Synopsis of the known
species of freshwater planarians of North America. Trans. Am. micr. Soc., 70 (2):
154-167.
, 1951b.—The Invertebrates: Platyhelminthes and Rhynchocoela. The acoelomate
bilateria. v. 2. vii + 572. (McGraw-Hill New York.)
GRASSE, P., 1961—Traite de Zoologie. T. iv (1). Plathelminthes, Mesozoaires,
Acanthocephales, Nemertiens. (P. de Beauchamp: Classe des Turbellaries: 37-212.)
944 pp. (Masson et Cie., Paris.)
VERRILL, A. H., 1892.—Marine planarians of New England. Trans. Conn. Acad. Arts &
Sci., iii (24): 459-520.
THE PLANTS GRAZED BY RED KANGAROOS, MEGALEIA RUFA
(DESMAREST), IN CENTRAL AUSTRALIA
G. M. CHIPPENDALE*
Forest Research Institute, Forestry and Timber Bureau, Canberra, A.C.T.
(Plates vi—viI1)
[Read 24th April, 1968]
Synopsis
A study of the diet of red kangaroos in an area of central Australia shows that
grass (75% to 99%) is the preferred fodder, with Hragrostis setifolia Nees being the
most important single species grazed. Forbs are significantly grazed only for short
periods in spring, and trees and shrubs are only grazed in small amounts with some
increases in summer.
During drought grazed Hragrostis setifolia is higher in moisture content than
ungrazed.
A pattern of grazing by kangaroos is suggested from the results obtained, but there
is evidence that a larger amount of grazing takes place in gilgais than-on the open
plain.
INTRODUCTION
Preliminary analysis of the diet of red kangaroos, Megaleia rufa
(Desmarest), in central Australia indicated that green herbage, and in
particular the grass Eragrostis setifolia Nees predominated (Chippendale,
1962). As Newsome (1965a, 6, c; 1966) has shown the supply of green
herbage to be vital to the red kangaroo in central Australia, breeding,
survival of young, and distribution and abundance depending on it, the plant
species grazed by kangaroos in different seasons were studied in detail. The
Burt Plain, an important drought refuge for kangaroos (Newsome, 1965q),
about 32 miles north of Alice Springs, was chosen as a study area.
HasBitat
The Burt Plain les between the Burt and Harry Creeks and is cut by
the Stuart Highway (Fig. 1). To the east of the highway, the open grassland
varies from Astrebla pectinata (Lindl.) F. Muell. ex Benth. communities
which were depauperate during the investigation to small water channels
and some gilgais carrying Hragrostis setifolia Nees and other grasses. To
the west of the highway, the grassland is predominantly Hragrostis setifolia
in and around gilgais, often mixed with Hragrostis xerophila Domin. The
gilgais had a particular assemblage of other species, including Swainsona
campylantha F. Muell., Psoralea cinerea Lindl., Neptunia dimorphantha
Domin., Alternanthera angustifolia R.Br., Abutilon malvifoliwm (Benth.)
J. M. Black, Oentipeda thespidioides F. Muell., Marsilea exarata A.Br., and
Portulaca oleracea lL. Occasional deeper gilgais were almost completely
dominated by Marsilea exarata. After rains, particularly in summer, annual
grasses grew on the plain and in the gilgais.
* Formerly Animal Industry Branch, Northern Territory Administration, Alice
Springs, N.T.
PROCEEDINGS OF THE LINNEAN Sociery oF NEw SouTH WALES, VoL. 93, Part 1
= G. M. CHIPPENDALE 99
The gilgais are marked depressions into which water runs after rain, and
at times (Plate v1) 50 or more points of rain would be sufficient to fill a
gilgai. The response of the vegetation to this water was spectacular (Plate
vil), with far more growth in the gilgai than on the surrounding open plain.
Tripogon loliiformis (F. Muell.) C. E. Hubbard occurred on the higher
parts of the plain after rain, and was most common in the heavy soil area
of Cotton Bush, Kochia aphylla R. Br.
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The grasslands were surrounded by Acacia aneura F. Muell. ex Benth.
woodland with sparse Hragrostis eriopoda Benth. present, and beyond the
mulga was the creek bank association with Eucalyptus camaldulensis Dehnh.,
Eremophila longifolia (R.Br.) F. Muell., Santalum lanceolatum R.Br.,
Dichanthium sericeum (R.Br.) A. Camus, and Chloris acicularis Lindl.
The rainfall on the Burt Plain, including for several months prior to
sampling, is shown in Table 1.
100 PLANTS GRAZED BY RED KANGAROOS
Cattle were present in small numbers for periods of several months
during the investigation.
MeEtTHODS
Samples from the stomach contents of kangaroos were taken at frequent
intervals from October, 1959 to October, 1961, except for five months in 1960,
the kangaroos being shot after dark, usually ‘when feeding on the plain west
of the highway.
TABLE |
Rainfall on the Burt Plain (points)
Year Jan. Feb. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dec.
1959 a Aaa” io) 8 155 —— 1 19.0 6 5 — 1 85 98 53
1969 .. .. 385 78 — 44 58 9 12 27 61 99 58 52
1961 .. so Se) 22 — 236 9 a
The samples, approximately 20% of the stomach contents, were washed
in a fine sieve, dried in the sun and stored in packets until examined. They
were later examined under a binocular microscope on an extension arm, using
the point frame method of Chamrad and Box (1964). A frame of crossed
threads sat squarely on the dried sample which was spread evenly in a
12” x 10” enamel dish. The microscope was used to identify the fragment
at each of the 100 points where threads crossed.
Artificial samples containing known percentages by weight were made
up by a Technical Assistant and examined by the author who had no
knowledge of the contents. Results of the test are shown in Table 2. It can
be seen that the accuracy for major constituents is better than for minor
constituents, and there is the possibility of missing some minor constituents.
TABLE 2
Results of test of point frame method, using artificial samples
Quantities (Percentage)
Ist Test 2nd Test 3rd Test 4th Test
Species
Point Point Point Point
Actual Frame Actual Frame Actual Frame Actual Frame
Teapeshe setifolia 95-0 95-0 50-0 52-0 80-0 82-75 97-0 99-25
EHnneapogon polyphyllus 1-0 3°0 —_— -- 2-0 5-75 —— =
Portulaca oleracea 2-0 0-75 10-0 5:2 2-0 0-75 — =
Euphorbia drummondi 0-5 0-5 — = 2-0 2-0 aod cia
Salsola kalz 1-0 0-75 — —- 2-0 0-75 1-0 —
Acacia aneura 0:5 a 10-0 8:75 2-0 0:75 1-0 0:5
Acacia victoriae — — 10-0 12-75 2-0 1-5 = =
‘Tribulus terrestris 3 — — 10-0 11-0 2-0 0-5 — ss
Atriplex elachophylla.. = — — 10-0 10-25 2-0 0-5 — —
Boerhavia diffusa — — = _— 2:0 2°75 — —
Indigofera domintt.. — — — —- 2-0 1-5 — —
In using the point frame method, four estimates were made for each
sample, two each by a Technical Assistant and the author, and the results
for each sample averaged.
Fragments were identified by reference to a standard set of plant
fragments collected on the Burt Plain, and by checking with authentic
specimens in the Herbarium of the Northern Territory, Alice Springs. Owing
to the difficulty of identification of grasses from minute fragments, mostly
G. M. CHIPPENDALE 10£
less than 1:0 mm. long, scores in the point frame were given as “grass”
initially, so that only a total grass percentage was obtained. Subsequently,
the sample was scanned under the microscope to identify the grass species
present and to assess relative dominance, but percentages for individual
species were not obtained. Grasses were sometimes identified by portions of
spikelets or spikes in the sample, and these in turn were related to leaf or
stem fragments associated in same sample. This information provided a
guide for identification in those samples in which leaves only were present.
When dealing with Hragrostis species, which were often present as leaf
fragments only, a means of separating these was sought. Examination of
the hairs on the inside of the convolute leaves iwas satisfactory; the angle
and arrangement of these hairs varied slightly in H. setifolia, E. xerophila
and EH. eriopoda (Plate viii), these being the main species available. Test
determination of fragments from herbarium specimens showed about 90%
positive accuracy, with the remainder being doubtful; there were no errors.
Fragments of leaves, stems, fruits and seeds in the samples aided the
identification of forb and topfeed species, but when there was doubt, specimens
were collected from the Burt Plain for comparison.
A number of kangaroo-proof exclosures each five metres square were
erected on the Burt Plain; two of these are pertinent to this project, each
being on a gilgai. Six transects across each exclosure were each continued
for a further five metres outside the fence on two sides; periodic measurements
were made of the basal length of any species on each transect. All measure-
ments were reduced to percentage density cover.
After results were known, the moisture content in grazed and ungraved
Eragrostis setifolia was measured by weight from monthly samples.
RESULTS
The stomach contents of red kangaroos on the Burt Plain contained some
green plant material at all times, though dried basal material predominated
in drier periods. This accords reasonably with the results reported by Griffiths
and Barker (1966) for Cunnamulla.
Table 3 shows that grass species dominated the diet, almost exclusively
doing so CATH the hot months, representing from 75% to 99% of the plants:
eaten.
Of 189 samples, 154 contained Hragrostis setifolia, and of the remainder,
9 contained Hragrostis xerophila and Tripogon loliiformis, 9 more contained
Eragrostis xerophila, 8 more contained Tripogon loliiformis, 1 contained
Astreba pectinata and Chloris acicularis, 7 contained mainly an assemblage
of annuals, and 1 contained unidentified grass species.
In dry times, such as from October, 1959 to January, 1960, Eragrostis
setifolia and Eragrostis axerophila were grazed by most animals; other species
found in the samples in this period were only as traces. With good summer
rain, Tripogon loliiformis became dominant in the diet in February, 1960.
After little effective rain during the winter, Hragrostis setifolia and
Enneapogon polypnyllus (Domin.) N. T. Burbidge were grazed by most
animals during August and September, 1960, but some short ephemeral
grasses also contributed to the diet. Moderate rain in September and
October caused the diet in early November, 1960 to be mainly Hragrostis
setifolia, Eragrostis xerophila, Tripogon loliiformis and Hnneapogon poly-
phyllus. By late November and until March, 1961, only dry vegetation was
available, and Eragrostis setifolia dominated the diet; Hragrostis xerophila
was commonly grazed in February.
PLANTS GRAZED BY RED KANGAROOS
102
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TaBLE 3—Continued
The quantity of grass and the species contained in samples from kangaroo stomach contents—Continued
Annuals
Perennials
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Range of
Quantity Quantity Relative
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Average
(%)
Number
of
Samples
Field Conditions
Sample
Period
1960
G. M. CHIPPENDALE
85 61-97
10
Some green
5 Sept.
97 95-99
10
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97
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GRAZED BY RED KANGAROOS
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G. M. CHIPPENDALE 105
Following the good rain in early April, 1961, Tripogon loliiformis
predominated in the diet, with Hragrostis setifolia. In May, 1961 with no
further rain, annual grass species contributed most to the diet, Dactyloctennum
radulans (R.Br.) Beauv., Tragus australianus S. T. Blake and Brachiaria
gilesii (Benth.) Chase predominating, with Iseilema membranacewm ‘(Lindl.)
Domin. and Enneapogon polyphyllus; seeding heads were common in the
samples at this time and Hragrostis setifolia and Tripogon loliiformis were
grazed by fewer animals. In June to August, 1961, Tripogon loliiformis and
Eragrostis setifolia were grazed by almost all animals with annual species
being grazed by slightly fewer animals. In October, 1961, the indication is
that Hragrostis setifolia had again become the main grass in the diet,
supported with some annual material.
Table 4 shows that forb species were mostly grazed in small percentages
and that the total forbs in the diet only exceeded 10% for short periods.
Calotis hispidula F. Muell., Marsilea exarata and Portulaca oleracea were
the most consistently recorded annual forbs, and Helipterum floribundum DC.
tended to become important during the months of August and September.
Several other species increased from trace amounts to at least 1% of the
diet for brief periods of from one month to several months, and these were
Helipterum charsleyae F. Muell. (included as “other Compositae”) in August,
September and October, Psoralea cinerea in September, 1960 and March, 1961,
Alternanthera angustifolia in January to March, 1961. A number of other
forbs were consistently recorded as being grazed in trace amounts usually
by less than half of the animals sampled at any period, and these were
Euphorbia drummondii Boiss., Neptunia dimorphantha, Indigofera dominii
Hj. Hichler, Abutilon malvifolium, and various Bassia species. Other forbs
were grazed in trace amounts during isolated periods by few animals.
Table 5 shows that woody species rarely comprised more than 1% of the
diet and then only for brief periods. Acacia aneura was the predominant tree
or shrub species grazed, being recorded in almost all months, but only grazed
by about half of the animals. When the amount of this species in the diet
increased from trace amounts to 3% in January, 1960 and to 2% in March,
1961, the number of animals grazing the tree also increased, as did the
number of tree species being grazed. Acacia tetragonophylla F. Muell. was
6% of the diet in October, 1961, after a long rainless period. Acacia sessiliceps
F. Muell. was grazed as 1% in January, 1961 and 2% in March, 1961 (listed
under other species). Other woody species were recorded as trace amounts
by few animals or in isolated periods.
Table 6 gives the percentage ground cover in and around two exclosures,
and it can be assumed that the cover inside would represent the available
fodder on the plain with no grazing while the cover outside reflects grazing
as well as an indication of actual available fodder on the plain. Table 6
considered with Tables 3-5 shows that on 16th November, 1960, there were
more forbs than Hragrostis spp. on the Burt Plain; yet grass comprised 91%
of the kangaroos’ diet. The diet remained about the same on 29th November,
1960, with available forbs now less than the grass. Grazing of forbs and
topfeed was increased on 29th March, 1961, when Eragrostis setifolia had
been markedly reduced outside the exclosure. Good rains from 10th to 19th
April, 1961, resulted in about equal amounts of forbs and grass being avail-
able inside, but herbage was less than half the amount of grass outside; the
diet at 27th April, 1961 was 995% grass with Hragrostis setifolia and
Tripogon loliiformis dominating. Forbs were again more plentiful than grass
on 26th May, 1961, but grass was still 99% of the diet, with the annual grasses
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4 0961
ic) = = ee Ae ee PCM C/E C(x ea eX a Gx SO) ONC pee =e Ol ‘00d 81-11
2 as 1/x ee | Sa XO ee) OU ee KK ei LTP N/M TS oS PI ‘190 61-81
= 6961
fy are
=|
° mh HO ® & FS & BHSWAaH BH Bw ACO Eb Ga FH GS BH SSTQ ay
3 =. 3. He soe 5 1 8 Sass TSI) GOS) SSS RS SR SUR aS
n sas z. S. S 5 | 8 Sees Soe : : : Bz. S$ Ss § § ee a = ss = SI S (%) setdureg powog
a i ae a xn a) ° aa S) SES e $ => = S 3 Ayyueng jo eidureg
SB Bee ees er eS % Be 3 5S requ
2) D ‘
pepiosea yorym ut sofdures jo roquimu/oseqyucoied osBI0AW
Se a
squajuoo yoowojs oowmbuny wouf saadups ua payfyuapr saweds quof fo sareguonb 04,7,
106
> FISvL
G. M. CHIPPENDALE 107
now being selected by more animals. A similar situation showed on 22nd
June, 1961, although the perennial grasses and some forbs were being grazed.
By 24th October, 1961, after no rain for six months, negligible amounts of
forbs were available on the plain and the perennial grasses were still the
dominant items in the Hel but forbs persed about 4% of the diet, with
topfeed 7%.
Figure 2 shows the amounts of moisture in samples of grazed and
ungrazed Hragrostis setifolia collected in and near gilgais on the Burt Plain
from November, 1963 to January, 1965. Rainfall is also shown. It can be
seen that during dry times the shorter grazed shoots contained more moisture
than the dry ungrazed grass. -
TABLE 5
The quantities of woody species identified in samples from kangaroo stomach contents
Average peccentage/number of samples in which
recorded
Sample Number Quantity
Period of (%) Acacia Acacia
Samples Acacia estro- tetra- Acacia Other
aneura phiolata gonophylla victoriae Species
1959
13-19 Oct. .. 14 1 1/11 x/1 x/3 x/3 x/2
11-18 Dec. .. 10 1 x/6 x/6 — > adil x/2
1960
8-14 Jan... 7 4 3/6 x/d x/3 x/1 x/2
21 Jan. .. 10 t x/7 x/1 — — x/1
4-12 Feb. 4 4 x/4 x/1 _ — —
19 Aug. 10 4 x/3 — x/1 — —
26 Aug. 10 3 x/5 x/1 — — =
5 Sept. 10 $ x/4 — — = =
3 Nov. 10 £ x/4 a _— — —
16 Nov 10 4 x/2 x/1 — = =
28 Nov 10 t x/6 = = = =
20 Dec 10 t x/2 — — = =
1961
16 Jan. 10 1 x/3 — _ x/1 1/4
22 Feb. 10 _ _ — —_ —
29 Mar. 10 6 2/6 — 1/3 1/3 2/7
27 Apr. 10 4 x/4 — — — —
25 May 10 t x/3 a = == =
22 June... 10 f x/2 — = = =
8) Ane S62 UO Ht x/3 — — x/1
24 Oct... 4 7 x/3 x/1 6/3 x/2 —
Total number of samples in which
species occurred (of 189) .. Ee 85 15 13 1] 18
x=less than 1%.
DISCUSSION AND CONCLUSIONS
Green grass is the preferred diet of the red kangaroo in central Australia.
Within a month, possibly within several weeks, of effective rain at any
time, the diet is almost exclusively grass. Observations at such times have
shown that Hragrostis setifolia and Tripogon loliiformis have soft green
regrowth, and this regrowth predominated in the stomach contents. Effective
rain in autumn caused annual grasses Tragus australianus, Dactyloctenium
radulans, Iseilema membranaceum, Brachiaria gilesii, and Enneapogon poly-
phyllus to be grazed by almost all of the animals sampled. These species
would probably be grazed after summer rain as they respond in summer.
108 PLANTS GRAZED BY RED KANGAROOS
It is notable that Hragrostis eriopoda, which is widely available to the.
kangaroos in areas of Acacia aneura, was not recorded as grazed. Similarly,
Astrebla pectinata which is available on the open plain, was rarely grazed.
With little effective rain in winter, the forb content of the diet increased
to its highest point of about 24% in August, 1960 and 16% in August, 1961.
At these times, the rapidly growing ephemeral Helipterum floribundum was
apparently selected, as high percentages were recorded. Perennial grasses
would be in poor condition at this time.
Portulaca oleracea, a succulent annual herb, was particularly selected,
although in comparatively small amounts, from November, 1960 to June, 1961.
This species, and Marsilea exarata, Neptunia dimorphantha, Indigofera
dominti, Psoralea cinerea, Alternanthera angustifolia and Abutilon malvi-
folium were usually grazed as an assemblage of plants, and as a group,
comprised from 1% to about 9% of the diet. As these grow in the gilgais
with Hragrostis setifolia, it indicates that most grazing takes place in finite
areas, the gilgais.
TABLE 6
Percentage cover in two exacloswres on the Burt Plain
Other Species, pre-
Eragrostis setifolia dominantly Forbs
Date
Inside Outside Inside Outside
16.11.1960 4-3 2-7 7-6 7:6
29.11.1960 8-6 6-4 7:8 5-0
20.12.1960 9-8 4-7 20-1 3:5
16. 1.1961 9-2 4-6 7-5 1-3
22. 2.1961 . 13-1 5-5 20-6 3°6
29. 3.1961 9-3 2-3 3-2 0-2
27. 4.1961 O19) 9-9 9-4 4-4
26. 5.1961 12-1 8-0 21-4 9-2
22. 6.1961 16-1 7-4 21-0 4-9
9. 8.1961 12-2 3-9 9-5 0-7
5. 9.1961 12-6 3-2 5-6 0-4
17.10.1961 11-4 3:8 3°8 0-2
27.11.1961 8-3 2-0 2-1 0-0
The extra water which runs into gilgais makes the grass more succulent,
and the moisture content of short shoots of Hragrostis setifolia which are
grazed ranges from 10% to 20% higher than that of taller ungrazed material
of the same species. Although water content was studied several years after
the main project, it is likely that similar conditions applied during the
project period.
Browse species are unimportant in the diet, for the main woody species
eaten, Mulga Acacia aneura, only comprised 2% to 8% of the diet during
January to March when extra roughage may be sought to balance the soft
grass after rain; it was grazed much less at other times. Other browse
species were mainly represented as traces. These small amounts would seem
consistent with small sporadic grazing in the Acacia aneura scrub where
the animals mainly rest during the day.
It is concluded that the short green shoots of the perennial grass
Neverfail, Hragrostis setifolia, in gilgais, was preferred by kangaroos on the
Burt Plain as the bulk of fodder all through the year. Tripogon loliiformis,
which is a perennial with a short growth period and is indicatively known as
Five Minute Grass, was also favoured in periods immediately after rain, and
G. M. CHIPPENDALE 109
probably as long as green leaf shoots are present. In late winter or early
spring, some annual herbage was mixed with the perennial grasses, and in
summer the animals select Portulaca oleracea with smaller quantities of other
annual herbs growing in gilgais. Several annual grasses are selected during
summer, and in mid-summer after rain small amounts of topfeed, notably
Acacia aneura, were eaten. Annual grasses again contributed during autumn.
From this pattern, and from observations, the kangaroo may not seriously
affect natural pastures, but certainly grazes the basal portions of perennial
grasses in dry times, and in a long drought, this could be serious. Trees are
rarely grazed, and no traces of bark or wood were seen in the samples. Field
observations confirm that the animals eat the soft regrowth of perennial
70
60
50
0
ial bee
Fs S
te)
< o
= =
aa 30 a
S
‘@
(ad
20
10
0
1963 1964 1965
Fig. 2. Moisture contents of samples of Hragrostis setifolia: -————— grazed, in
gilgais; ---—-—— not grazed, near gilgais.
grasses in gilgais and also graze to a lesser degree other plants which may
be green. Similarly, tall grasses are not grazed; it was common to see mature
stands of Hragrostis spp. on the open plain as ungrazed, while at the same
time, tracks, faecal pellets, and signs of grazing in gilgais were common. An
explanation seems to be that more or less constant grazing in the gilgais kept
the grasses actively growing, and it seems possible that the congregation of
fzecal matter at these points together with the run-on effect of rainfall may
give added recovery power to the grass.
As the perennial grass Hragrostis setifolia is the most important single
species in the diet of the kangaroo in central Australia, it is possible that
this grass in gilgais, and the red kangaroo, may at times form an inter-
dependent relationship.
110 PLANTS GRAZED BY RED KANGAROOS
Acknowledgements
I acknowledge gratefully the great amount of work done in this project
by Mr. D. J. Nelson, Technical Officer, Animal Industry Branch, Northern
Territory Administration, Alice Springs, N.T. Mr. Nelson shot the kangaroos,
prepared the samples, made some examinations with the point frame, and
also erected the exclosures; he also constructed the point frame and offered
suggestions on the manuscript. I am grateful to Mr. R. Swinbourne, Botanic
Garden, Adelaide, South Australia, formerly of the Animal Industry Branch
at Alice Springs, for assistance in the field and in the herbarium. The
determinations of moisture content in Fig. 2 were made by Dr. B. Seibert,
Chemist, of the Animal Industry Branch, Alice Springs. Figures 1 and 2
were finally drawn by Miss Marjorie Hall, Forest Research Institute, Forestry
and Timber Bureau, Canberra, A.C.T., from draft figures prepared by Mr.
Nelson. Plates vi-vi1 were taken by the author, and Mr. A. Edwards of the
Forest Research Institute, Canberra, took Plate v111.
I appreciated criticism of my manuscript by Dr. A. E. Newsome, Division
of Wildlife, C.S.I-R.O., Canberra, A.C.T.
References
CuHaAmrapD, A. D., and Box, T. W., 1964.—A point frame for sampling rumen contents.
J. wildl. Mgmt., 28: 473-477.
CHIPPENDALE, G. M., 1962.—Botanical. examination of kangaroo stomach contents and
cattle rumen contents. Aust. J. Sci., 25: 21-2.
GRIFFITHS, M., and BArKeEr, R., 1966.—The plants eaten by sheep and by kangaroos
grazing together in a paddock in south-western Queensland. O.8.1.R.0. wildl. Res.,
11: 145-67.
NeEwsomgE, A. E., 1965a.—The abundance of red kangaroos, Megaleia rufa (Desmarest),
in central Australia. Aust. J. Zool., 13: 269-87.
, 1965b.—The distribution of red kangaroos, Megaleia rufa (Desmarest), about
sources of persistent food.and water in central Australia. Aust. J. Zool., 13: 289-99.
, 1965c.—Reproduction in natural populations of the red kangaroo, Megaleia
rufa (Desmarest), in central Australia. Aust. J. Zool., 13: 735-59.
————., 1966.—The influence of food on breeding in the red kangaroo in central
Australia. O.S.1.R.0. wildl. Res., 11: 187-96.
EXPLANATION OF PLATES
PLATE vi. An exclosure on a gilgai on the Burt Plain, central Australia, 19th
April, 1961.
PLATE vil. The same exclosure and gilgai, 27th April, 1961.
PLATE vit. Patterns of hairs on inner surface of the convolute leaves of: Top left,
Eragrostis xerophila from herbarium specimen R. Winkworth 617, right NT313; Centre
left, Hragrostis setifolia from NT9011, right NT9012; Bottom left, Hragrostis eriopoda
from M. Lazarides 5189, right N. Burbidge and M. Gray 4544.
PLATE VI
8, Part 1
¢
Proc. Linn. Soc. N.S.W., Vol.
T96T
‘
lady W161
‘
UIeTq Jang ot
UO IBS[IS B
uo dINSO[OUS UY
Proc. Linn. Soc. N.S.W., Vol. 98, Part 1
PLATE VII
T96T
‘
THdy wg ‘Ureld
yng oy} uo
IBs[Is B UO sINSsO[DUD UV
iW]
Proc. Linn. Soe. N.S.W., Vol. 98, Part 1 PLATE VIII
eis Ry Male il
oe naily
lie om
Se ne
a M 6: pee ties
hee. aa x EE ‘e 46
a Ging DPR Peer res Se
gee Eee adh a ‘ oe]
Pattern of hairs on leaves of HWragrostis species.
THE LOWER AND MIDDLE PALAEOZOIC STRATIGRAPHY AND
SEDIMENTARY TECTONICS OF THE SOFALA-HILL END—
EUCHAREENA REGION, N.S.W.
G. H. PackKHAM
(Plates 1x—xI1)
[Read 24th April, 1968]
Synopsis
Ordovician andesitic volcanics are the oldest rocks in this region of about 1,000
square miles. They are overlain about Sofala in the east of the region, by Silurian
limestones and shales, rhyolitic volcanics, subgreywackes, dacitic tuffs and further
subgreywackes, and west near Euchareena by rhyolitic volcanics, limestone and shale.
The source of the sediments lay to the west and south-west except for the highest
subgreywacke unit in the Sofala sequence which was derived from the underlying
formations as a result of the uplift of a structure to the east (the Capertee Geanticline).
Dacitic vuleanism on this geanticline in the early Devonian provided directly or
indirectly, much of the detritus comprising the greywackes and tuffs of the Crudine
Group and the Merrions Tuff which were deposited in the Hill End Trough in the
Sofala-Hill End area. In the Euchareena district more restricted andesitic and dacitic
vulcanism occurred at this time. During the later Lower Devonian and perhaps part
of the Middle Devonian, greywacke sedimentation took place in the Hill End Trough
while limestones and shales were deposited in the Limekilns area. Near Huchareena
limestones, shales and lithic calecarenites were being laid down. The youngest rocks
deposited in the region prior to the Upper Devonian are acid tuffs overlying the lime-
stone-shale sequence at Limekilns perhaps unconformably. The region is folded into
a broad synclinorium which has a culmination in the vicinity of Hill End and is
overthrust to the east near Sofala. The rocks of the central part are strongly cleaved
and bélong to the green schist metamorphic facies.
INTRODUCTION
The region of over a thousand square miles, described in this paper
contains folded strata ranging in age from Ordovician to Devonian. Within
the region there are three palaeogeographic and tectonic entities (Figure 1),
which have been discussed in previous publications (Packham, 1960, 1962,
1968). They are from east to west, the Capertee Geanticline, the Hill End
Trough and the Molong Geanticline. This paper presents the stratigraphic
evidence on which the recognition of these features was originally based.
Within the Capertee Geanticline, which came into existence at about the
end of the Silurian, the sediments are normally uncleaved and only moderately
folded. Limestones occur in the Lower Devonian and indicate that shallow
water environments occurred in that part of the succession at least. The
geanticline has as its western boundary the Wiagdon Thrust, a structure
which is now known to extend from north of Mudgee to south of Yetholme,
a distance of about 100 miles.
The Hill End Trough which occupies most of the region under discussion
contains greywacke-type sediments throughout the exposed Silurian and
Devonian section. Folding is moderate to strong, with slaty cleavage developed
everywhere in the finer sediments and in the coarser rocks in the central
part of the trough. The cleavage fans on a regional scale, dipping to the
PROCEEDINGS OF THE LINNEAN Society or New SoutH WALES, VoL. 93, Part 1
112 PALAEOZOIC STRATIGRAPHY AND SEDIMENTARY TECTONICS
west in the eastern part and east in the west. It is approximately vertical
in the vicinity of Hill End. The highest grade of metamorphism reached is
in the central part of the trough where biotite is developed over a width
of about 10 miles across the strike. On the western side of the trough, the
greywacke sequence passes into fossiliferous sediments on the eastern flank
of the Molong Geanticline where limestones occur in the Ordovician, Silurian
and Devonian Systems.
The region is one in which outcrop is generally good. It is essentially a
plateau with an elevation of about 3,200 feet in the east and 2,500 feet in
the west, strongly dissected by the Turon and Macquarie Rivers and their
tributaries. Exposures along the rivers and the major tributaries are excellent.
The description of the stratigraphy is divided into three sections (a)
the Sofala-Limekilns district of the eastern side of the Hill End Trough and
the western edge of the Capertee Geanticline (6b) the Hill End district, in
the centre of the trough and (c) the Euchareena district on the western side
of the trough and the eastern side of the Molong Geanticline.
Rock specimens mentioned in the text are in the rock collection of the
Department of Geology and Geophysics, University of Sydney.
THe STRATIGRAPHY OF THE SoraLA—-LIMEKILNS DIstTrRIictT
In the Sofala district there is an essentially conformable sequence of
strata, ranging in age from Upper Ordovician at the base to possibly Middle
Devonian at the top. The highest unit in the Limekilns area, the Winburn
Tuff, may be unconformable on the Limekilns Group. The successions in the
Sofala-Turondale area and the Limekilns area are as follows (in descending
order) :
Sofala-Turondale Limekilns _
Winburn Tuff 2000+’
Cunningham Formation 2800’ Limekilns Group 2500’
Merrions Tuff 2000’ Merrions Tuff 1500’
Crudine Group 3700’ Crudine Group 2500’
Cookman Formation 1500’ Cookman Formation 600’
Chesleigh Formation 3500’ Chesleigh Formation 3500’
Bell’s Ck. Voleanies 1500’ ——.
Tanwarra Shale 250’ Tanwarra Shale 0-250’
Sofala Voleanics te Sofala Voleanics 7000+’
The Limekilns sequence is east of the Wiagdon Thrust and the Sofala sequence
is west of this fault in the Hill End Trough.
SoraLta VOLCANICS
The type section of the Sofala Volcanies is exposed along the Turon
River, commencing one and a half miles west of Sofala, extending ten miles
to the east where the formation is thrust over Upper Devonian sandstones.
No underlying formation has been recognized so that the base of the Sofala
Voleanics cannot be defined. In the type section, approximately 7,000 feet
of strata consisting dominantly of clastic and pyroclastic detritus with a
small proportion of lavas, are exposed. Occasional horizons of chert represent
concentrations of pelagic elements.
Type section. In the lower part of the section medium to fine-grained
sediments are dominant, typically in beds a foot thick, displaying graded
bedding, convolute bedding and slump structures. Fine-grained sediments in
G. H. PACKHAM iLilss
the section are very dark grey to black in colour and have a sub-conchoidal
fracture. Three types of coarser sediments occur in the section. There are
breccias containing a small amount of matrix, composed almost entirely of
fragments of rocks of the same lithology as the fine-grained sediments in the
section (Plate rx, fig. 1). The second type consists of an abundant fine-
grained matrix including rounded cobbles and boulders of andesite and silt-
stone blocks up to a foot or so in diameter. The third kind, composed of
angular blocks of intermediate lavas occurs sporadically in the lower part
of the type section but is abundant in the upper part. About the middle of
|: = |Silurian
|:::]Ordovician [* *]Granite
| - |Devonian
N
720s \(1
J
i
\
Bul DIUV qe
Molong = —+*—— Hill End Trough —_—_——+ Capertee
Geanticline Geanticline
Figure 1.General regional map showing distributions of Ordovician, Silurian and
Devonian Systems and major structures and tectonic units mentioned in the text.
the sequence (about 24 miles east of Sofala) there is a chert horizon several
hundred feet thick which extends north and south of the Turon River for
several miles.
The upper part of the formation consists almost entirely of tuffs,
abundant andesite breccia and some andesites. West of Sofala these are
transgressed by dykes and other minor intrusions of rocks of lamprophyre
affinities, apparently basic differentiates of the andesites. The increase in
the coarseness of the volcanics and the presence of the related intrusives
suggest that the vent (or vents) responsible for the deposition of the upper
part of the section may have been close at hand.
Regional distribution and variation. In the upper part of the sequence,
limestone blocks occur in breccias in Big Oaky Creek just east of Wattle
Flat, and a limestone lens occurs one and a half miles north-north-west of
Wattle Flat. The limestones may have resulted from the rapid building up
of voleanic debris in the upper part of the section. Apart from this, the
formation maintains the general characteristics of the type section throughout
its outcrop.
H
114 PALAEOZOIC STRATIGRAPHY AND SEDIMENTARY TECTONICS
Except in close proximity to the Wiagdon Thrust where cleavage is
developed, the rocks of the Sofala Voleanics are massive and indurated.
Even the fine-grained sediments have little tendency to split along bedding
planes. Small movements along joints are observable in almost every outcrop.
Petrography. Apparently most of. the lavas of the Sofala Volcanics were
originally andesitic. They contain phenocrysts of plagioclase, augite and/or
hornblende. Occasional lavas have groundmasses containing small plagioclase
laths but in most the groundmass is very fine and even grained. These latter
may have been originally glassy. The plagioclase in all the thin-sections
which were examined is albite. The ferromagnesian minerals are fresh in
most rocks, except in the area north-west of Sofala. Two types of hornblende
occur in the relatively unaltered rocks; green hornblende with a pleochroic
scheme, Z = olive green, Y = yellow green, and X = pale yellow with ZAC
about 24° and brown with Z/\C about 18°. The green amphibole often shows
signs of resorption and the brown amphibole occasionally contains inclusions
of pyroxene. The pyroxenes are pale green in thin section and have a moderate
2V and an extinction of about 45°.
The lamprophyres which occur to the west of Sofala are petrographically
very similar to the andesites except for the absence of felspar phenocrysts.
An almost identical association has been described from the Wellington
District by Basnett (1942). The Sofala lamprophyres are amphibole-rich,
but at least some of this replaces pyroxenes as is evidenced by the typical
pyroxene form of some of the amphibole crystals and the presence of rare
irregularly shaped cores of pyroxene. The replacing mineral has a Z/\C of
20° and X = colourless, Y and Z = pale green. This mineral also replaces
darker amphibole in which X = yellow, Y = brown and Z = brownish green
and Z/\C is about 27°. The groundmasses of these rocks are highly altered,
now consisting of albite, pale green amphibole and -epidote with calcite
abundant in some.
The volcanic breccias are composed almost entirely of fragments of
various types of andesites. There is no sign of rounding even in the larger
fragments. The matrix consists of small rock fragments and grains of felspar
and pyroxene. Near the top of the succession where there is considerable
alteration, it is not clear whether those rocks containing angular blocks up
to several inches across are actually lavas including fragments of andesite
similar to those described by Basnett and Colditz (1946) from rocks of
comparable age at. Wellington, or whether they are actually breccias. The
presence of numbers of clastic fragments in some localities suggests an
original sedimentary origin for the rocks. The matrix in which the fragments
now occur is composed of an interlocking mass of albite, epidote and pale
green amphibole. In the lower part of the sequence, relatively unrecrystal-
lized breccias consist of fragments of lavas (generally a variety of types),
pale green pyroxene a little felspar (albite) in a matrix of chlorite, calcite
and epidote.
Like the breccias, the rocks of intermediate grainsize consist almost
entirely of volcanic material. Except in the upper part of the formation
where the bedding is obscure, the sandy sediments are greywackes. They are
poorly sorted. Felspar (albite) is the dominant mineral, the grains are either
idiomorphic or broken crystal fragments. Green and brown hornblende and
augite are present. A little quartz occurs, this shows signs of magmatic
corrosion and is hence probably volcanic in origin. Rock fragments are rare,
those present are grains of the groundmass of andesite. The matrix, which
usually comprises some 20 percent of the rock, is composed of chlorite, albite
and a little calcite.
j
'
G. H. PACKHAM 115
The finer grained rocks have a mineralogy similar to that of the arenites.
The finest of these sediments are chert-like in hand specimen and are difficult
to distinguish from the occasional radiolarian cherts which occur throughout
the sequence. The radiolarian rocks themselves, consist of poorly preserved
radiolaria, making up roughly 10 percent of the rock, in a fine-grained ground-
mass of material showing weak double refraction and a refractive index higher
than canada balsam—presumably quartz. Large numbers of minute black
granules, possibly carbonaceous, and a few percent of albite and chlorite
grains of silt size are scattered through the fine groundmass of the radiolarian
rocks. Several series of quartz veins must have been deposited soon after
deposition of the sediment since where fragments of the radiolarian rocks
are incorporated in intraformational breccias, they sometimes contain quartz
veins predating their incorporation (Plate rx, fig. 1). If the fragments were
derived from older formations it is most unlikely that in such a sediment
all the fragments would be angular and of precisely the same lithology as
the finer sediments in the Sofala Volcanics. Deformation seen in some of the
early formed veins in the radiolarian rocks is probably the result of the
continuation of compaction of the sediment after the formation of veins.
Depositional environment and direction of origin. The association of
sedimentary structures in the lower part of the Sofala Volcanics is as follows:
graded-bedding, convolute-bedding and slump structures. Cross bedding is
absent. This association together with the presence of graptolites and radio-
larian cherts suggests a deep water environment of deposition.
The beds of coarse sediment with rounded andesitic cobbles and boulders
with a large proportion of matrix are not simple pyroclastic sediments. The
rounding of the andesite cobbles indicates some transportation before final
deposition. Slumping or some form of mass movement seems to be the only
way to explain the association of these sediments with what are almost
certainly deep water sediments. Contemporaneous movements have resulted
in the. breaking up of some of the fine-grained sediments of the sequence
resulting in breccias composed almost entirely of fragments of those rocks.
Towards the top of the formation the appearance of limestone in blocks
and lenses suggests that there was some shallowing which may be due to
orogenic movement, local volcanic accumulation or filling of the entire basin.
Slump structures observed in the type section, a short distance upstream
from Eaglehawk Gully indicate movement from jwest to east. Thus an easterly
sloping sea floor can be inferred.
Age. Poorly preserved graptolites have been found about the middle of
the Sofala Volcanics in Eaglehawk Gully. The graptolites appear to be
Glyptograptus teretiusculus which is found in uppermost Darriwillian and
Gisbornian strata. If this identification can be relied upon, then it is prob-
able that much of the Ordovician is represented by the volcanics since the
graptolites occur a little below the middle of the exposed section of the
formation. The top of the formation pre-dates the Tanwarra Shale, the age
of which, from the contained graptolites and other lines of evidence, is
probably Upper Llandovery or early Wenlock. The upper possible limit of
the Sofala Voleanics is within the Lower Silurian.
TANWARRA SHALE
This formation rests, with a slight break, on top of the Sofala Volcanics.
The junction of the two formations is obscured in many localities by over-
thrust faulting on the Wiagdon Thrust. The base of the Tanwarra Shale is
marked by a conglomerate or sandstone resting on top of the Sofala Volcanics.
116 PALAEOZOIC STRATIGRAPHY AND SEDIMENTARY TECTONICS
This basal member is composed almost entirely of material derived from the
erosion of the andesites, together with some fossil detritus.
Type section. The type section of the formation is in Portion 516 of the
Parish of Sofala, in the headwaters of Spring Creek and Bell’s Creek, south
of Mount Tanwarra after which the formation is named. This section is
250 feet thick. The basal conglomerate is of the order of forty feet in thickness
and contains limestone pebbles with “Halysites” in addition to andesitic
detritus. Above, is an impure limestone about twenty feet thick. This lime-
stone consists of alternating bands of fossil material and sandy calcareous
shale. The larger coral fossils are preserved in their living environment or
have been transported only a short distance. The fauna is as follows:
Phaulactis shearsbyi, Mucophyllum liliiforme, Heliolites daintreei, Acantho-
halysites cf. gamboolicus, Alveolites sp., Atrypa sp., ef. Barrandina sp. and
Encrinurus sp. Above this horizon there are a few feet of shales and sandy
shales. The shales which make up the remainder of the formation are green,
grey and black in colour, all are poorly bedded and contain occasional fossil
remains, mostly poorly preserved small brachiopods. Ten feet above the
limestone there are sparse graptolite remains. The forms present are: Mono-
graptus cf. paradoxzus and M. priodon.
The top of the formation is recognised by the appearance of bands of
acid tuff marking the beginning of the Bell’s Creek Volcanics.
Regional distribution and variation. North of the type section, the
formation is considerably sheared. The stratigraphy of the formation in this
area is not clear. It is possible that north of the Turon River some of the
rocks mapped as Tanwarra Shale may actually be Chesleigh subgreywacke.
The sediments are rolled out to such an extent that in many exposures
bedding has been completely destroyed and replaced by a crude alternation
of silty and shaly material parallel to the cleavage.
To the south of the type section there is a small outcrop of fossiliferous
marl a mile to the west of the top of Wiaedon Hill which may be a south-
wards continuation of the formation; from this Hncrinurus sp. was collected.
At Wiagdon Hill on the Bathurst Road, the formation is represented only
by a thin limestone and some shales; this is on the overthrust side of the
Wiagdon Thrust. In the creeks to the south-east along the strike the forma-
tion is missing, apparently faulted out. Underlying the Bell’s Creek Volcanics
three miles south of Wiagdon Hill there is a sequence of shales representing
a continuation of the Tanwarra Shale.
There is only one known occurrence of the Tanwarra Shale on the eastern
side of the fault. This is in the head of the Spring Creek in Portion 245 of
the Parish of Sofala near the western border of Portion 253, three and a
quarter miles east of Wattle Flat. The succession is similar to that of the
type section four and a half miles to the west, except that graptolites have
not been found at this second locality. The fauna includes: ef. Phaulactis
shearsbyi, Mucophyllum crateroides, Tryplasma sp., Spirinella caecistriata
and Hnerinurus sp. Fossils are considerably more abundant and_ better
preserved than in the type section.
In the sections east of the Wiagdon Thrust where the Tanwarra Shale
is absent the Sofala Voleanics are followed by a breccia composed of andesitic
detritus. This is succeeded by the Chesleigh Formation without the inter-
vention of the Bell’s Creek Volcanics.
Age. The age of the formation is clearly Silurian. If the Monogr aptus cf.
paradorus is in fact M. paradoxus then the age is almost certainly Upper
Llandovery; if not, then the age could be anywhere from the base of the
pie er
G. H. PACKHAM a7,
Upper Llandovery to half way up the Wenlock. The presence of “Halysites”
in the shelly fauna is also in agreement with an age in the lower half of the
Silurian.
BELL’S CREEK VOLCANICS
The Bell’s Creek Volcanics is a succession of rhyolitic tuffs and lavas
which conformably overlies the Tanwarra Shale. The formation varies
considerably in thickness along the strike and is confined to the western
side of the Wiagdon Thrust.
Type section. The type section, 1,500 feet thick, is exposed in the valleys
of Bell’s Creek and Jew’s Creek, two miles south:west of Sofala. This is the
maximum thickness known for the formation. The basal part consists
dominantly of tuffs, succeeded by rhyolite and then by another tuffaceous
unit. Overlying is the Chesleigh Formation, the lower part of which is non-
voleanic. The lavas of the Bell’s Creek Volcanics form a high ridge to the
west of Bells Creek while the underlying tuffs and a dolerite sill occupy
the floor and the eastern side of the valley. The tuffs at the top of the
formation occupy the small valley of Middle Creek to the west of the ridge
of rhyolites.
Regional distribution and variation. North of the Turon River the
formation is considerably diminished in width as a result of thrust faulting.
It is possible that the contact between the Bell’s Creek Voleanics and the
Tanwarra Shale north of the Turon River might be a tectonic one.
The Bell’s Creek Voleanics outcrop in the Wiagdon Hill road section as
tuffs, slates and chert-like rocks. Half mile to the south-east very poorly
preserved graptolites occur in a cherty shale in the formation. They are
straight monograptids but no further identification is possible. Further south
in the vicinity of Cheshire’s Creek, lavas are once more abundant. The
volcanics can be traced for eighteen miles along the strike but to the east
of the Wiagdon Thrust the formation is unknown. This is unexpected in
view of the considerable extension of the volcanics along the strike and it
is the first indication of .a significantly different stratigraphic succession on
either side of the thrust.
Petrography. The rhyolites are pale green or buff coloured rocks with
a subconchoidal fracture. Flow-banding is only rarely visible in hand
specimen. In thin section these rocks have either a uniform groundmass or
irregular flow structure. Phenocrysts, which are not abundant, are dominantly
quartz and orthoclase, with minor amounts of albite (about Abg;). Idiomorphic
phenocrysts are rare, mostly they are corroded or fractured. Ferromagnesian
minerals, present in only small quantities, are confined to the groundmass.
They are either biotite, pleochroic from yellow to very dark brown, or chlorite,
which is apparently the alteration product of the biotite. Epidote and
clinozoisite are present in some rocks occurring as small individual crystals
scattered through the rock and as aggregates up to 1 mm. across. The lavas
are cut by quartz veins some of which also contain clinozoisite or epidote.
The groundmass shows a good deal of textural variation even within the
one section; it varies from a very fine-grained to a coarse-grained mosaic of
interlocking quartz and felspar grains.
The tuffaceous lithologies in the formation range from coarse breccias
to extremely fine-grained rocks with a flinty appearance. Their mineralogy
is similar to that of the rhyolites with which they are associated. The medium-
grained tuffs are composed of broken quartz and felspar grains, biotite and
devitrified glass shards, together with a little apatite. The finer tuffs are
118 PALAEOZOIC STRATIGRAPHY AND SEDIMENTARY TECTONICS
banded with some evidence of graded-bedding on a small scale. The extremely
fine-grained tuffs are generally pale green flinty rocks, without bedding in
hand specimen or with broadly developed irregular banding. In thin-section
these are seen to consist of an interlocking mass of quartz and felspar with
some calcite, chlorite and epidote.
CHESLEIGH FORMATION
The Chesleigh Formation takes its name from the property “Chesleigh”
(three miles west of Sofala) on which it outcrops as a prominent strike ridge
and conformably overlies the Bell’s Creek Voleanics. It is non-voleanie in
the lower part of its section, thus the base can be defined as the top of the
last tuff horizon.
Type section. The formation is 3,500 feet thick in this section which is
that exposed along the Turon River in Portions 48, 47 and the eastern part
of Portion 46 of the Parish of Waterbeach. The lowest unit, a massive sub-
sreywacke, 2,000 feet thick, outcrops on the south bank of the Turon River
and forms a large cliff in which bedding is poorly defined. Overlying this
is 500 feet of interbedded slate and subgreywacke. The beds vary from a
few inches to ten feet in thickness. In the top thousand feet of the formation,
the amount of felspar increases, subgreywackes pass into greywackes and
there are some interbedded crystal tuff horizons in which the beds are up
to twenty feet thick. The uppermost beds are thinly bedded fine-grained
felspathic siltstones and slates. These siltstones which are green and red
are followed by the first of the quartzrich sediments of the Cookman
Formation.
Regional distribution and variation. In spite of the large size of out-
crops along the Turon River, there is very little fresh rock exposed in the
type section. Two miles to the north, on Two Mile Creek, the bedding
characteristics of the lower massive subgreywacke can be seen. Bedding
planes are spaced six to eight inches apart and each bed is to some extent
graded. The clarity of this grading is obscured by the small size range—
from fine sandstone to siltstone. The bedding features are further obscured
as a result of tectonic movement which has taken place within the formation.
This movement is particularly evident on the banks of Crudine Creek between
Two Mile Creek and the Turon River, where slaty cleavage is strongly
developed in interbedded slates and subgreywacke. To the south of the type
section, the subgreywacke lithology is dominant and the upper tuffaceous
part of the formation consists only of slates and fine tuffs with a cherty
appearance.
The Chesleigh Formation occurs on the eastern side of the Wiagdon
Thrust with lithology and thickness similar to those of the type section, but
differing in that cherty rocks are common in the upper part of the formation.
In this region, the Chesleigh Formation rests directly on the Sofala Voleanics
except in the headwaters of Spring Creek where the Tanwarra Shale occurs.
The contact between the Tanwarra Shale and the Chesleigh Formation in
this area has not been observed.
The absence of the Bell’s Creek Volecanies and the restriction of the
Tanwarra Shale on the eastern side of the Wiagdon Thrust, suggests that
the base of the Chesleigh Formation in that area is an erosional surface.
The extension of the missing formations along the strike is such that they
might be expected to continue across the strike as well. However, it has
been already pointed out that the Bell’s Creek Voleanics vary considerably
from place to place. Similarly, since the Tanwarra Shale was, at least in
~ G. H. PACKHAM 119
part, laid down in a shallow water environment, it may perhaps have not
been deposited in some localities. Breccia composed of andesitic material
underlying the Chesleigh Formation east of Wattle Flat, mapped as part
of the Sofala Volcanics, might be the time equivalent of the Tanwarra Shale.
Even if there is no erosional break below the Chesleigh Formation, the
significant feature is that the Wiagdon Thrust brings into juxtaposition two
significantly different stratigraphic sequences. Differences between the sections
on either side of the thrust are maintained in most of the formations over-
lying the Chesleigh Formation.
Petrography. In the type section the subgreywacke (Plate 1x, fig. 2)
which makes up the lower part of the formation contains fifty to seventy
percent of quartz. Most of the grains show some sign of rounding, though
this is more evident in the largest grains (0-5 mm.). The sorting is poor,
all gradations exist between the largest grains and the finest material
constituting the matrix. This matrix makes up twenty percent or more of
the rock. Apart from quartz silt, the matrix is composed of chlorite and pale
green, weakly pleochroic mica. A small proportion of rock fragments is
present; these fragments are of two types. Firstly, shales consisting of a
fine-grained aggregate of quartz, chlorite and sericite and in some cases a
little carbonate. Secondly, there are fragments composed of interlocking
quartz grains with small amounts of chlorite either included in the grains
or concentrated in their margins. These latter rock fragments are probably
fragments of thermally metamorphosed sediments. Felspar is absent. In the
matrix there are occasional grains of muscovite. The heavy mineral
assemblage is varied and includes: tourmaline (variety of shapes, sizes and
colours), zircon, sphene, apatite and rutile.
In the upper part of the formation, the rocks which have been called
ereywackes differ from the underlying subgreywackes only in the addition
of felspar and a corresponding decrease in the percentage of quartz. Orthoclase
is more abundant than plagioclase (albite).
The tuffs of the Chesleigh Formation are massive with grainsizes ranging
from fine sand to breccias. Their interpretation in thin-section is difficult.
There is a good deal of secondary albite in the matrix and replacing
orthoclase, as well as in veins. Quartz and carbonate are the abundant
secondary minerals in some specimens while in others the assemblage is
albite, quartz and epidote. The detrital minerals are quartz, orthoclase,
plagioclase (now albite) and chiorite. In some of the coarser rocks fragments
of acid porphyritic rocks occur.
The lithology of the formation on the eastern side of the Wiagdon
Thrust is very similar to that of the type section. Some of the subgreywackes
however, contain a small proportion of felspar (orthoclase and albite), and
have a greenish colour in hand specimen instead of the normal grey colour.
The variation of colour is attributable to the presence of additional chlorite.
Depositional environment and origin of the detritus. A variety of sedi-
mentary structures indicative of a deep-water depositional environment, are
present in the Chesleigh Formation. They include graded bedding, load casts
(Plate 1x, fig. 8), small slumps, small-scale cross-bedding, and flute-casts.
The last three structures have yielded evidence indicating the direction of
slope of the seafloor at the time of deposition.
Small-scale slump structures occur near the top of the Chesleigh
Formation in the type section and in Dam Creek, in fine-grained siltstone
beds four to eight inches thick. The direction of movement which has been
determined by measuring the inclination of a number of slump fold axes in
each locality, is consistently towards the east-north-east.
120 PALABEOZOIC STRATIGRAPHY AND SEDIMENTARY TECTONICS
A similar direction of movement is indicated by flute-casts beautifully
exposed on the bases of overturned beds two miles north of Cheshire’s Creek
and a locality on Cheshire’s Creek half a mile east of the Wattle Flat—Peel
road. At the first locality the direction of movement is corroborated by the
dip of small-scale cross-bedding in a similar direction.
The detritus in the lower part of the formation is apparently first or
second cycle detritus derived from a granitic terrain. The low proportion of
felspar and the presence of sedimentary rock fragments suggest the former
rather than the latter. It is known that there were folding movements in
the Tasman Geosyncline (the Benambran Orogeny) roughly at the end of the
Ordovician Period and continuing into the Silurian (see Packham, 1967q).
The Ordovician sediments exposed in the southern highlands of New South
Wales by these movements jwere slates, shales and subgreywackes which
were derived from granitic rocks directly or indirectly forming an ideal
source for the detritus for the Chesleigh Formation.
In the upper part of the Chesleigh Formation acid volcanic material is
common but detritus of the type which makes up the lower part of the
formation is still present. '
Fauna. No fossils have been found in this formation.
CookKMAN FORMATION
The Cookman Formation which consists of quartz-rich greywackes,
occasional grits and conglomerates interbedded with slates overlies the
Chesleigh Formation. Its base is recognized by the first appearance of the
characteristic light-coloured quartzite-like subgreywacke and sublabile grey-
wacke.
Type section. The type section of the Cookman Formation, one thousand
five hundred feet thick, is in the eastern half of Portion 46 of the Parish
of Waterbeach on the south bank of the Turon River immediately west of
the type section of the Chesleigh Formation. The formation takes its name
from the Cookman Range which is the strike ridge that the formation makes
to the north of the Turon River. The type section consists dominantly of
slates with interbedded fine-to-medium-grained sandstones and only a few
beds of coarse sandstones and conglomerates. Most of the sandy beds, several
inches to a foot thick, display no obvious internal structure, but the beds of
the order of three feet thick are graded-bedded. Frequently, casts of worm-
tracks appear on the under surfaces of the sandy beds. At the top of the
formation there is a thick slate which is overlain by the lowest coarse-
grained beds of the Crudine Group. ;
Regional distribution and variation. The Cookman Formation maintains
the features of the type section along its strike. It can also be recognized
east of the Wiagdon Thrust overlying the Chesleigh Formation. In Cheshire’s
Creek in the southern part of Portion 119 (Parish of Wiagdon), the formation
is similar in lithology to the type section but only six hundred feet thick.
It can be traced north from Cheshire’s Creek for some distance but is lost
in the area of poor outcrops south-east of Wattle Flat.
Petrography. The typical quartz-rich medium-grained arenite (grains
up to 0-6 mm.) is moderately sorted with only a small percentage of matrix
(Plate 1x, fig. 4). The quartz grains vary from rounded to angular.
Identifiable mineral inclusions are rare, though rutile, apatite and yellow
tourmaline have been observed. Flakes of muscovite up to half a millimetre
long are scattered through the rock. The matrix is composed of quartz silt
with white mica and some chlorite. In the heavy mineral assemblage zircon
a
G. H. PACKHAM 12]
is the most abundant mineral, both idiomorphic and rounded grains of com-
parable size are present; the rounded ones are far more abundant. Grey-
green, brown and blue tourmalines have been noted; almost all show signs
of rounding. Occasional grains of rutile occur. The most significant, yet
least abundant mineral present in the heavy mineral assemblage is horn-
blende, pleochroic from yellow-green to grass-green. The finer--grained sand-
stones have a smaller percentage of felspar than the medium-grained ones
and the sorting is better; the number of large grains is reduced with
decreasing grain-size though the percentage of matrix remains much the same.
The coarser sediments are more varied in their features. Micrometric analyses
of some of the rocks’ of the formation are given in Table 1. Felspar and rock
fragments become increasingly abundant in the coarser rocks. Both orthoclase
and plagioclase (albite) are present; occasionally the plagioclase crystals
have rows of inclusions arranged in zones indicating that there were once
composition differences within the crystals. Some orthoclase is partially
replaced by albite. The rock fragments are mostly finegrained, dominantly
TABLE 1
Micrometric analyses of Arenites from the Cookman Formation
A B C D
Quartz .. oe ae ok: $l
Rock fragments By or 8
Muscovite a At ie 1
Chlorite grains .. l
Felspar (Or and Ab) 1
Sericite-quartz matrix .. or! 10
Calcite .. ie N: re —
|Selloas
A. $387. Maximum grain-size 0:-6mm. One mile south of type section.
B. TS 109. Maximum grain-size 1-5 mm. Type section.
C. TS 110. -Maximum grain-size 3 mm. Type section.
D. TS 118. Maximum grain-size 5 mm. Type section.
fragments of the groundmasses of acid porphyritic rocks. Occasional large
aggregates of quartz and felspar are present, possibly derived from granitic
rocks. Rare detrital grains of microcline endorse this view. A little detrital
carbonate is present both as crystalline rock fragments and as patches in
the matrices of these coarser sediments.
The slates of this formation are composed of white mica, chlorite, and
variable proportions of quartz silt. The abundance of mica is responsible
for their pale grey colour in hand specimen.
Depositional environment and source of the sediment. East of the
Wiagdon Thrust flute casts are well exposed on the bases of the overturned
beds. The lobes of the flute-casts are narrower and less rounded than those
in the Chesleigh Formation. Good exposures of these structures which are
found on Cheshire’s Creek and three miles north of it, indicate a direction
of flow a little south of east. This is the opposite to that in the Chesleigh
Formation. Thus, the change in direction of slope of the sea-floor implies
the uplift of an area to the east. This eastern land mass incidentally, had
an extremely important role in the later history of the region. The petro-
graphy of the Cookman Formation suggests that a variety of rocks were
eroded off this newly exposed area, including acid volcanics (indicated by
rock fragments and corroded quartzes), sediments which were quartz-rich
(rounded quartzes, heavy mineral assemblage with abundant rounded zircon
and less abundant rounded tourmaline), possibly granite (microcline, quartz-
felspar aggregates and possibly idiomorphic zircons) and basic igneous rocks
1D PALAROZOIC STRATIGRAPHY AND SEDIMENTARY TECTONICS
(amphibole in the heavy mineral assemblage). All of these source rocks can
be accounted for in the underlying formations except for the possible granite.
The material derived from the acid volcanics may have come from the erosion
of the volcanics at the top of the Chesleigh Formation or the Bell’s Creek
Voleanics. The sediments which gave rise to the majority of the heavy mineral
assemblage belonged, in all probability, to the Chesleigh Formation which
would in addition, have provided the greater part of the quartz of the
Cookman Formation.
The increase in the proportion of unstable rock and mineral fragments
in the coarser fractions of the sediments of the Cookman Formation may be
explained by considering the weathering characteristics of the suggested
source rocks. The subgreywackes of the Chesleigh Formation which at this
stage would be still unconsolidated, and therefore erode by disaggregation
of the grains yielding small particles. The tuffs may erode by a similar
process but the coarser types would give rise to the larger particles. The
lavas would be eroded by the normal processes of weathering and abrasion
and give rise to particles of a variety of sizes. Thus, if a sediment is com-
posed of larger grains, then it must, of necessity, be derived from a source
other than the subgreywackes of the Chesleigh Formation. The medium to _
fine-grained sediments of the Cookman Formation would, on the other hand,
be composed dominantly of material derived from those subgreywackes. The
presence of amphibole can be related to the Sofala Voleanics, mainly on the
grounds that there are no other formations in the sequence known to contain
amphibole. The location and age of the granite which was suggested as a
possible contributor to the detritus is completely unknown.
The environment of deposition of the Cookman Formation seems to have
been a deep-water one. There is some evidence of life on the sea-floor in the
presence of worm tracks. The greater elongation of the flute casts in Cookman
Formation suggests that the slope may have been greater in this area during
the deposition of the Chesleigh Formation.
Age. No fauna has been found in this formation but fragmentary plant
material was collected on the top of the high hill formed by the formation,
three quarters of a mile south of the Turon River. These fragments are
sometimes Y-shaped and are similar to some of the plant material found in
the Silurian and Devonian in Victoria. Their age is certainly no older than
early Ludlow.
CRUDINE GrRouP
The Crudine Group is so called because much of the valley of Crudine
Creek is cut into these sediments. There is a sharp change from the quartz-
rich sediments of the Cookman Formation to the massive tuff and breccia
at the base of the Crudine Group. The top of the group is the base of the
very thick and massive Merrions Tuff. This is a distinctive boundary since
the upper formation of the Crudine Group is one which consists of grey-
wackes and slates and contains no tuff horizons.
The Crudine Group is several thousand feet thick and contains a variety
of rock types, all typical of eugeosynclinal sedimentary associations; pyro-
clastics which range from coarse breccias, down to fine-grained chert-like
tuffs and the normal clastic sediments which include breccias, conglomerates,
sands, silts and muds all of the greywacke suite, but no chemical and organic
sediments. It is difficult to distinguish between tuffaceous rocks and grey-
wackes in some instances because the dominant source of the sediments is
voleanic.
= G. H. PACKHAM i2R3
Type section. The group consists of two formations; the Turondale
Formation below and the Waterbeach Formations above. The aggregate
thickness of the type sections of the two formations is 3,700 feet.
a. Turondale Formation. The type section of this formation, which takes
its name from the nearby village of Turondale, is on the south bank of the
Turon River in Portion 46 of the Parish of Waterbeach overlying the type
section of the Cookman Formation. The Turondale Formation is 2,020 feet
in this section and consists of three units (Figure 2).
The lowest division of the formation, 830 feet thick, is composed almost
entirely of tuffaceous material. Only 120 feet are not derived directly from
voleanic sources, these are slates and fine-grained sandstones. The basal bed
800'44, == LEGEND
Greywacke Massive buffs
Tuffaceous breccia Banded tuffs
1900/4"
1800"
1200°
300 1700’
1100"
200 1600’
1000
100’
Figure 2. Type section of the Turondale Formation measured on the south bank of
the Turon River in Por. 46, Parish Waterbeach. Greywacke beds less than 2 feet thick
are shown as thin horizontal lines.
of the formation is a coarse conglomerate in which the matrix is at first
fine-grained and makes up a large proportion of the rock but towards the
top of the bed the amount of matrix decreases to about half of the rock
and becomes sandy. The pebbles in the conglomerate are mainly of acid
voleanic rocks with some of limestone, shale and tuff. Limestone pebbles and
boulders are more abundant in the lower part of the bed. Angular shale
blocks contemporaneously derived, are often very large; the largest seen
had exposed dimensions of ten feet by three feet. The conglomerate passes
upwards into volcanic breccia. This breccia and the succeeding tuff, which
are typical of the tuffaceous sediments in the Crudine Group and the Merrions
124 PALAEOZOIC STRATIGRAPHY AND SEDIMENTARY TECTONICS
Tuff, consist of quartz, felspar, chlorite and epidote. They are hard and
massive, pale green rocks with only poorly developed bedding planes with
beds of the order of tens of feet thick (Figures 2, 6, 8). Graded bedding
is commonly developed over the entire thickness of the bed. The coarser
phases often contain a small percentage of shale fragments and rounded
felsite pebbles but most of the fragments are irregular, flattened chloritic
bodies. These latter generally have inclusions of quartz and felspar, which
are often embayed. Epidote and zircon are also present (Plate rx, fig. 3).
The constant appearance of these bodies in sediments rich in volcanic debris
in the region suggests that they have a volcanic origin. A possible explanation
is that they were originally angular glassy fragments which have been
subsequently converted to chlorite, possibly having passed through an inter-
mediate stage of palagonite. This type of phenomenon has been described
by Tyrrell and Peacock (1926) and Raw (1948). Other products of this
process were, in all probability, subsequently converted to epidote. The very
fine-grained rocks at the top of this basal bed have ill-defined bedding and
a conchoidal fracture and are often translucent on the thin edges of fresh
specimens. Their colour ranges from pale green through greenish-grey to
dark grey. Sometimes they have spots several] millimetres in diameter
scattered through them in indefinite bands; these are apparently the result
of local concentrations of chlorite and epidote. Above these tuffaceous
sediments there are interbedded fine-grained greywackes and slates which
are followed by more massive tuffaceous rocks.
The second unit, 580 feet thick, is dominantly slate and silt with bands
of greywacke and a few tuffaceous beds. The tendency of these greywacke
beds to occur in groups is a striking feature which is repeated at higher
levels in the Waterbeach Formation and, to a lesser extent, in the Cunningham
Formation. The greywacke beds differ in appearance from the tuffaceous
beds below in a number of ways. The greywackes are thinner, mostly one
to three feet thick, but ranging from several inches to ten feet. Shale pebbles
are the most common of the larger rock fragments. Graded bedding is common
but by no means universal. When present there is a gradation from grey-
wacke to shale, whereas the tuffs, as described above, grade into cherty rocks.
The slates interbedded with the greywackes are in units several inches thick,
the bedding being marked by a thin silt band which occasionally shows
suggestions of small-scale cross-bedding and graded bedding.
The upper part of the Turondale Formation, 610 feet thick in the type
section, contains a proportion of tuffaceous material which is reminiscent
of the lower part of the Turondale Formation, but the thicknesses of the
beds are closer to those of the middle unit. In hand specimen, these rocks
are Seen to contain few rock fragments, chloritic patches are rare and so are
shale fragments. There is little indication of graded bedding, where present
the gradation is into shale in some beds and cherty rocks in others. In the
fine to medium sand size-range some beds have alternating pale green and
dark green bands, one to three inches thick, resulting from variation of the
proportion of chlorite in the rock.
b. Waterbeach Formation. This formation takes its name from the
parish in which the type section is defined. This section, 1,690 feet thick, is
on the south bank of the Turon River in Portions 69, 74 and 49, a little over
a mile west of the type section of the Turondale Formation. In the Water-
beach Formation, shaly rocks are far more abundant than sandy ones and
tuffaceous rocks are absent. The base of the formation is the top of the
last tuffaceous band of the Turondale Formation. There is a marked change
in the characteristics of the sedimentation at this point. The greywackes are
G. H. PACKHAM 125
in distinct groups at 300, 800, 1,100 and 1,300 feet above the base of the
formation. With the exception of shale pebble bands which occasionally occur
at the bases of the thicker greywackes, conglomeratic rocks are absent from
the section. A generalized stratigraphic column is given in Figure 3.
The sandy zones may represent times of greater local tectonic activity
which aided erosion and hence provided coarser detritus, or alternatively,
the control could be purely climatic, since periods of greater rainfall would
have the same effect. Eustatic changes provide a third possible mechanism.
LEGEND
Greywacke
penne silbs
1600’
1500"
14.00’
1300‘
500°
1200°
400°
1100°
1000’
DOO a= seee
200°
100
800°
Figure 3. Type section of the Waterbeach Formation measured on the south bank
of the Turon River in Portions 41, 74 and 69, Parish Waterbeach. Greywacke beds
less than 2 feet thick are shown as thin horizontal lines.
It is unlikely that they represent volcanic outburts because there are no tuff
horizons associated with them. Further, the petrography of the sediments
indicates that material has been only partly derived from a volcanic source.
The majority of beds of greywacke and siltstones (over four inches thick)
are graded-bedded. Some of the thinner silt beds are graded-bedded, passing
up gradually into the overlying slate. Others form overall graded units with
oscillations of grainsize within the unit. Still others have no grading, they
have distinct silt bands with irregular bedding, small scale cross-bedding, or
cross bedded ripple marks. In all types the silts have sharp bases frequently
126 PALAEOZOIC STRATIGRAPHY AND SEDIMENTARY TECTONICS
infilling animal trails on the top of the underlying slate unit. The silty units
are generally thinner than the slate between them, the distance from the
base of one silt unit to the next varying from two to six inches (Plate x,
fig. 1). The thinnest beds of greywackes are the most abundant and there is
a progressive decline in the frequency of occurrence of beds with increasing
thickness. The thickest bed observed has a thickness of twenty-four feet. In
the composite graded beds which occur occasionally, there is no parting
along the junction of the coarse greywacke with the underlying fine grey-
wacke. Occasionally, the lower unit has been dragged up into the upper unit
during the deposition of the latter. Most of the graded beds maintain almost
the same grain-size over the lower three quarters of their thickness then
decrease gradually in grain-size in the remainder of the bed. This upper
part of the bed which is normally fine greywacke or siltstone, generally has
bedding planes spaced about a quarter of an inch apart. Only very
infrequently is there a gradation into slate. It is possible that the beds
which have no graded bedding observable in the field are beds in which the
upper finer grained part of the bed has not been deposited.
The greywackes often contain angular fragments of shales which have
been derived, in all probability, from contemporaneously deposited sediments
by the erosive action of sediment charged turbidity currents. Normally, shale
fragments included in the greywackes are four to five inches long and an inch
or so thick. The largest shale fragment observed in this formation has
exposed dimensions of five feet long and five inches thick. It is in a grey-
wacke bed only three and a half feet thick. Shale fragments are the only
fragments coarser than sand size found in this section of the formation.
Groove casts, flute-casts and load-casts occur on the bases of some of
the greywacke beds in this formation, but their frequency is difficult to
determine because the bases of the beds are infrequently exposed. Only a
few slumps have been observed in this section, all of them are in the thinly
bedded slate-siltstone part of the sequence. Each slump involves a foot or
less of strata.
Regional variation of the Crudine Group. The Turondale Formation
changes its character both across and along the strike. The most important
new feature is the appearance of conglomeratic phases in the upper unit.
These conglomerates are found north and south of the type section but have
their greatest development three to five miles south of the Turon River where
they are several hundred feet thick. In this conglomerate there is at least
60 percent of medium-grained greywacke matrix keeping the pebbles well
Separated (Plate x, fig. 2). The pebbles are varied in type; they include
quartzite, quartz porphyry, limestone and shaly rocks. Quartzite and quartz
porphyry are rounded, their size varying from less than an inch to six to
eight inches along their greatest diameter. The limestone and shale fragments
have been deformed along with the matrix during the folding of the succession
and as a result they are flattened into lenses in the cleavage plane. By far
the largest block of limestone observed measures 80 feet by 30 feet; it occurs
in Portion 45 of the Parish of Waterbeach. The occurrence of this block
of pure limestone containing tabulate corals, in a coarse conglomerate and
its general association with graded-bedded rocks, together with the absence
of any other limestone lenses and the remains of shelly fossils from the
associated sediments all support the contention that it is a derived block.
It is most unlikely that large pebbles and cobbles were carried by a turbidity
current; it is more likely that they were carried by a mud-flow, or deposited
by sliding or slumping. Further south, conglomerates are less frequent. In
spite of this variation the Turondale Formation is dominated throughout by
a ees
= G. H. PACKHAM WA
detritus of volcanic origin. The Waterbeach Formation is more constant in
its characters. The only significant change in features is that to the north
of the type section commencing a mile from the Turon River, a thick grey-
wacke unit containing some conglomeratic phases is developed near the base
of the formation.
The section east of the Wiagdon Thrust. The section here is thinner
(2,500 feet), the tuffaceous beds are more distinct, graded-bedding does occur
(Plate x, fig. 3), but is not common. The coarser beds are better sorted than
in the Turon River section. Further to the east, on the eastern limb of the
Jesse Syncline near. the top of the group, there are fragmentary brachiopods
in mudstone; this is the only locality in the Crudine Group where invertebrate
fossils other than those occurring in limestone boulders have been found.
One horizon of coarse conglomerate consisting of large quartz porphyry
cobbles in a matrix of sand derived from similar material is exposed on
Cheshire’s Creek.
Direction of source of the sediments. The lithological variation described
above leads one to conclude that the sediments in the eastern side of the
Wiagdon Thrust are closer to their source than the rocks to the west. This
view is backed up by the evidence provided by the sedimentary structures
which occur in the Crudine Group. In the type section of the Waterbeach
Formation between 1,100 feet and 1,350 feet above the base of the formation,
there are a number of structures, flute-casts and small-scale cross-bedded
current-ripples, which indicate a direction of sediment flow towards a little
north of west. Three and a half miles north of the type section near the
top of the Waterbeach Formation and five miles south of the type section
at much the same horizon there are occurrences of small-scale cross-bedding.
The former indicates a direction of flow a little south of west and the latter
towards the west. Similar structures occur high in the Crudine Group east
of the Wiagdon Thrust one mile west of Limekilns Public School site. At this
locality, the structures indicate a slope towards the west-north-west.
Thus, at all the localities where there is any evidence of slope of the
sea-floor, the general direction of slope indicated is towards the west. So the
same palaeogeographic conditions must have existed during the deposition
of the Crudine Group as existed at the time when the Cookman Formation
was deposited.
Petrography and mode of deposition of the sediments. The sediments of
the Crudine Group have few constituents. The grains of sand and silt size
are: quartz, felspar, igneous rock fragments and occasional grains of calcite.
These are set in a matrix of chlorite and epidote with zircon, pyrite and.
apatite as minor constituents. Micrometric analyses of these rocks are set out
in Tables 2 and 3.
The tuffaceous rocks of the Turondale Formation differ somewhat from
the greywackes of the Waterbeach Formation. The greywackes contain a
larger proportion of rock fragments, a larger proportion of quartz relative
to felspar and a smaller proportion of epidote in the matrix. These differences
are brought out in Figure 4. Figure 5 illustrates the general resemblances
between the sediments. All these arenites fall into the field of labile grey-
wackes (Packham, 1954) and are thus distinct from the sediments of the
underlying Cookman Formation which fall into the fields of subgreyiwackes
and sublabile greywackes.
The most striking difference between the formations is in the nature of
the bedding. The frequency of occurrence of greywacke beds of a given
thickness in the Waterbeach Formation is roughly inversely proportional to
128 PALAEOZOIC STRATIGRAPHY AND SEDIMENTARY TECTONICS
the thickness of the bed. The tuffaceous rocks of the Turondale Formation
do not show this feature at all; if anything, the thicker beds are more
common than the thinner ones. This is also true of the Merrions Tuff which
overlies the Waterbeach Formation. Figure 6 illustrates this difference
clearly.
The problem which now arises is that of determining the mode of
deposition of the tuffs. The tuffaceous rocks have normal graded-bedded
TABLE 2
Micrometric analyses of Tuffaceous Arenites from the Turondale Formation
A B Cc D
Quartz .. bs se wie 20 8 12 27
Felspar.. oj 5a ey 34 32 31 27
Igneous rock fragments 56 9 6 2 8
Chlorite patches aie ag 6 oo 3 2
Chlorite matrix ae Si dl 54 52 36
All specimens are from the type section of the formation.
A. TS 55. 250 feet above the base of the formation.
Be TS 48% 875 feet above the base of the formation.
C. TS 45. 1,300 feet above the base of the formation.
D. TS 39. 1,470 feet above the base of the formation.
sediments overlying, underlying and interbedded with them, and it is difficult
to maintain that emergence has taken place so that the tuffs could be
deposited in shallow water. In any case the beds lack any characters which
would positively distinguish them as shallow water sediments. The Merrions
Tuff extends across the strike for twenty miles and a greater distance along
the strike. Such a distribution is out of keeping with the sediments being
simply the result of explosive volcanic activity. It is clearly impossible for
TABLE 3
Micrometric analyses of Greywackes from the Waterbeach Formation
A B C D E FE
Quartz aM as aR 15 16 28 17 13 14
Felspar af a e 21 1 9 19 17 4
Igneous rock fragments .. 23 30 27 20 30 37
Sedimentary rock fragments — 1 4 2 = 4
Chlorite-silt matrix a 41 4] 32 42 40 4]
All specimens are from the type section of the formation.
A. TS 34. 280 feet above the base of the formation.
B. TS 38. 440 feet above the base of the formation.
C. TS 24a. 1,100 feet above the base of the formation.
. TS 20. 1,350 feet above the base of the formation.
. TS 18. 1,350 feet above the base of the formation.
. TS 16. 1,600 feet above the base of the formation.
Aa
material of sand size and larger to be hurled such distances. In any case,
massive beds of the thickness found could not have been deposited by such
a process.
Textural similarity of the tuffaceous rocks and the normal greywackes
(Tables 2, 3; figs 4,5; Plate 1x; figs 5, 6, 7) and the negative arguments cited
in the previous paragraph lead me to the conclusion that the tuffaceous rocks
were deposited by the same sort of mechanism as the greywackes, ie., by
turbidity currents or as mudflows. A terrestrial mudflow, the Osceola mudflow,
i G. H. PACKHAM 129
described by Crandell and Waldron (1956) has a number of features
resembling the tuffaceous beds of the Turondale Formation and the Merrions
Tuff. The Osceola mudflow has a maximum thickness of 350 feet where it
has been restricted in a narrow gorge, but the thickness decreases to less
than 20 feet on the plain below. The formation is coarsely graded-bedded
(the concentration of boulders being much higher at the base than ‘the top).
The proportion of clay and silt in the sample analysed by Crandell and
Waldron is 39:6 percent. This accords reasonably well with the results I
have obtained for the sediments of the Turondale Formation and the Merrions
Tuff (Tables 2, 4) where the range is from 31 to 54 percent. Apart from the
Quart3
aT$39
5 1S8
alces PT42 S8l
e xX TSél
ats45 81220" 5e
Felspar Rock Fragments
x Cunningham Formation @ Merrions Tuff
e Waterbeach Formation A Turondale Formation
+ Cookman Formation
Figure 4. Plot of composition of the sand fractions of arenites listed in Tables 1-7.
greater proportion of boulders in the Osceola mudflow, its similarity. to the
tuffaceous rocks of the Turondale Formation and the Merrions Tuff is a
close one. The lateral change of upper part of the Turondale Formation
from tuffaceous rocks to greywacke conglomerate can be explained by
assuming that the mudflow involved had not moved a great distance and
that the variation of composition represents the influence of varied source
materials.
If the tuffaceous sediments are, in fact, mudflows, then strictly speaking,
it is not advisable to call such sediments “tuffs”, because although they may
have been intially deposited near a volcano, they were later transported to
their present location. I have, however, retained the term “tuff” to distinguish
I
130 PALAEOZOIC STRATIGRAPHY AND SEDIMENTARY TECTONICS
these sediments from the greywackes which, though clearly derived from
voleanic rocks for the most part, show no petrographic evidence that they
were contemporaneous with the vulcanism.
If alternating intervals of vulcanism and quiescence are postulated, the
differences between the two types of sediment may be explained. During the
volcanic episodes, large amounts of detritus of sand grade were made avail-
able, hence there is a preponderance of sandy detritus over finer material.
In the quiescent episode normal erosion of the terrain took place yielding
large amounts of silt and clay. Weathering of felspar in this terrain was
Matrix
4TSi10 Bee
TS20 gts34
EDO rere S14
TEGO Gl SUSI SUSE
TS240“TS55
4+7TS109 "Tse oo
x
+ TSI1&
X SBI O21
x SO
Felspar + Rock
Fragments
Quarta
x Cunningham Formation @ Merrions Tuff
© Waterbeach Formation _& Turondale Formation
+ Cookman Formation
Figure 5. Plot of composition of the sand fractions of arenites listed in Tables 1-7.
probably responsible for the increase of the quartz relative to felspar in the
Waterbeach Formation. Erosion of lavas extruded in the vicinity of the vents
could explain the larger proportion of rock fragments.
Comparison of the frequency of occurrence of beds in this succession with
those obtained by Potter and Siever (1955), for the Lower Pennsylvanian
and Upper Chester sandstones (orthoquartzites) in Illinois (Figure 7), clearly
brings out the difference between deep-water geosynclinal deposition and
shallow-water deposition. In the geosynclinal environment the frequency of
the beds in the one to four feet class is very high (70 percent) and drops
rapidly, while in the shallow water environment the frequency of sandstones
rises to a maximum of 20 percent in the four to eight feet class, and then
falls slowly with increasing thickness.
p G. H. PACKHAM 131
The quartz grains in the tuffaceous sediments in the Turondale Forma-
tion are mostly angular but occasional grains have bipyramidal form, gen-
erally with slightly rounded angles. Corrosion embayments are common and
are filled with fine grained material which constituted the groundmass of a
lava or filled with chlorite. The largest quartz grains are two millimetres
in diameter and it is these which are most frequently bipyramidal and
embayed. Plagioclase (about Abo;) is more abundant than orthoclase. The
grains of albite are always cloudy occasionally with inclusion of epidote and
almost always surrounded by epidote which has developed in the chlorite
matrix. The albite grains have the same maximum grainsize as the quartz
grains and, like them, sometimes have corrosion embayments filled with
Greywackes of Greywackes of
Cunningham Formation Waterbeach Formation
60 60
RESO Apo
Cc Cc
8 sy
2 2
5 40 5 40
a a
\ \
pp 30 # 30
c c
: :
¢ 20 x 20
3
2s =
we ive
10 10
fo) fe)
O 2 4 8 16 32 64 128 Oneealienicns2
Thickness of beds — feet Thickness of beds~ feeb
Tuffs of Turondale Formation
and Merrions Tuff
&
°
3
(eo)
O 2 4 8 16 32 64 128 256
Thickness of beds — Feet
Figure 6. Frequency distribution of bed thicknesses in the type sections of the
Merrions Tuff, the Cunningham and Waterbeach Formations.
Frequency - percent
chlorite. The albite grains are more often idiomorphic than the quartz grains.
The morphological characteristics of the orthoclase grains are similar to
those of the albite. In a number of thin-sections from the Turondale Forma-
tion the orthoclase grains have patchy replacement by albite similar to those
in the upper part of the Chesleigh Formation. Glass shards, now replaced
by albite, are common in some of the tuffaceous sediments; these offer posi-
tive evidence of a volcanic origin for at least some of the detritus. Very
rarely, the sediments contain grains of brown amphibole. In the darker
rocks, chlorite and epidote make up the entire matrix, in the lighter ones,
albite is present also. Rock fragments are not common in the tuffaceous
rocks, when they are present, they are acid lavas similar in mineralogy to
the tuffaceous rocks.
132 PALAEOZOIC STRATIGRAPHY AND SEDIMENTARY TECTONICS
The petrography of these tuffaceous rocks confirms the pyroclastic
origin of the detritus in a number of ways; glass shards are present. Quartz
and felspar grains show signs of corrosion (for some of the quartz grains
this is certainly magmatic, in others it might have been post-depositional).
Chlorite patches, probably derived from glassy rock fragments, are common.
There is no evidence from the petrography of the sediments or the associated
conglomeratic pebbies that the quartz and felspar grains were derived from
a plutonic source. The small proportion of rock fragments in the sandy
fraction precludes the possibility of the sediments being the result of the
weathering and erosion of acid lavas and porphyries.
Lower Pennsylvanian Upper Chester
Orthoquart3ites of Orthoquartzibes of
Ilinois Illinois
a)
C ic
9 9
2 30 2 30
5) 9
a. a
{ 20 (
x 5,20
9 0
9 5
S\i0 8 10
or om
9 RS)
ic © in ©
O 4 8 16 32 64 128 256 O 4 B 16 32 64 128
Thickness of beds-feeb Thickness of beds~Feet
Greywackes and Tuffs
of the Cunningham Formation
Merrions Tuff and Crudine Group
i
® db a ®
fo} °o Oo fe)
=3
Frequency - percent
b&b
fe)
3
(0)
O 4 8 16 32 64 128 256512
Thickness of beds -feeb
Figure 7. Frequency distribution of bed thicknesses in shallow-water sandstones in
Illinois and greywackes of the Hill End Trough.
The greywackes of the Waterbeach Formation are finer-grained than the
typical tuffaceous sediments of the Turondale Formation. The quartz and
felspar (albite with minor orthoclase) grains are smaller and more angular
in the greywackes. Igneous rock fragments are far more abundant in the
greywackes than in the tuffaceous rocks. Fragments over a millimetre in
diameter are normally rounded, smaller ones are angular. Shale fragments
are the most abundant sedimentary rock fragments, they are frequently
large and almost always angular. Limestone fragments are seen in thin-
section. The matrix is of chlorite, quartz, felspar and some scattered grains
of epidote. The proportion of epidote is much smaller than in the tuffaceous
rocks. The proportions of the various constituents in. the coarse and medium-
grained greywackes do not vary greatly with grainsize in any one bed. The
= G. H. PACKHAM 133
micrometric analyses of TS 18 and 19 (from the top and bottom respectively
of a graded-bedded greywacke, Plate 1x, figs 7, 6) illustrate this. The
proportion of rock fragments decreases noticeably only in the finer sand-
stones and siltstones.
The mineralogy of the siltstones and slates throughout much of the
Crudine Group is closely related to that of the coarser grained rocks asso-
ciated with them. The ratio of quartz to felspar is similar and so far as
can be determined in thin-section, the remainder of the rock is chlorite.
The distinctive appearance of fine-grained chert-like sediments of the
Turondale Formation results from their having a smaller proportion of
chlorite than the slates, considerable quantities of epidote and interlocking
erains.
Fauna and age. The only fossils found in the Crudine Group are ones
which have been transported from their living environment. Near the top
of the group approximately 5 miles north of Limekilns in Portion 107 of
the Parish of Jesse a mudstone has yielded a brachiopod fauna from which
Wright (1966) has identified Dolerorthis, sp., Skenidioides sp., [sorthis sp.,
Schizophoria sp., Plectadonta sp., Notanoplia sp., Schelwienella ? sp.,
Hospirifer sp., Ivanothyris ? sp., Spinatrypa sp., Lissatrypa lenticulata and
the coral Pleurodictyum. In the type section of the Turondale Formation,
limestone pebbles at the base of the formation contain Tryplasma sp.,
Favosites sp. and Pentamerid brachiopods. Near the top of the same section
very poorly preserved plant remains have been found. Three miles to the
south, in the upper part of the formation, limestone blocks contain Favosites
sp. close to F’. richardsi. The age indicated by the brachiopod fauna at the
top of the group is Lower Devonian.
Merrions Turr
This formation is extremely widespread and has proved of great value
in elucidating the structure and succession of the region. The base of the
formation is the base of the first tuff horizon after the greywacke succession
of the Waterbeach Formation. Its top is the top of the highest tuff bed in
the Hill End Trough succession where Merrions Tuff is overlain by the slates
and greywackes of the Cunningham Formation. In the vicinity of Limekilns,
the Merrions Tuff is overlain by shales of the lower part of the Limekilns
Group. In all sections, the Merrions Tuff consists dominantly of medium
sand-grade in massive beds of the order of tens of feet thick, these beds
often show grading on a large scale, from breccias at the base to fine chert-
like rocks at the top. Dacitic lava flows occur but make up only small
proportion of the formation.
The large-scale graded-bedded units commence with a basal breccia
which contains large chloritic bodies (Plate x, fig. 4) of the type described
in the account of the Turondale Formation, sometimes angular blocks of
shale and less frequently, rounded pebbles of quartz porphyries and lime-
stone. The matrix is composed of coarse sand-grade mineral fragments
(albite, orthoclase, quartz with interstitial chlorite and epidote). This is
succeeded by progressively finer detritus with an alternation of light and
dark bands several inches thick, the colour banding apparently resulting from
variation of the chlorite content. Sometimes just below the banded rocks
there is a zone which has a mottled appearance. This too, results from an
uneven distribution of chlorite through the rock. The banded rocks pass
upwards into finer tuffs, poorly bedded, light in colour and often with
indefinite slump structures. These rocks have a chert-like appearance and
134 PALAEOZOIC STRATIGRAPHY AND SEDIMENTARY TECTONICS
are frequently spotted. They have been described in the discussion of the
Turondale Formation.
Type section. The type section (Figure 8) of this formation is along the
south bank of the Turon River in Portion 41 of the Parish of Waterbeach,
overlying the type section of the Waterbeach Formation. The name of the
LEGEND
== Interbedded silts aT :
& slates NAY Dacite
Tuffaceous breccia Massive buffs
Greywacke Fine buffs
Banded tuffs
i000’
400’
Figure 8. Type section of the Merrions Tuff measured on the south bank of the
Turon River in Portion 31, Parish Waterbeach.
formation is derived from Merrions Trig. Station south of the type section
on a high ridge formed by the outcrop of the formation. The measured
thickness of the type section is 2,020 feet, this order of thickness is typical
of the formation. The formation can be divided into four units in this
43 G. H. PACKHAM 135
section. The lowest 475 feet of the formation contains four thick tuff beds.
The lower two commence with coarse tuff and pass upwards into banded
tuff which is followed abruptly by coarse tuff. The upper two tuffaceous
beds pass into indurated shaly rocks with characters similar to those of the
Waterbeach Formation. Interbedded with the fine-grained sediments at about
470. feet above the base of the formation there are two greywacke beds with
all the characters of the typical greywackes of the Waterbeach Formation
except that they contain a larger proportion of quartz and felspar and a
correspondingly smaller proportion of rock fragments.
- The next 600. feet of the formation is almost entirely coarse tuff. Two
exceptionally thick beds occur. The lower one is over 150 feet thick and the
upper one is over 200 feet thick. Shale blocks included in the base of the
lower one are up to two feet across while the upper one has large chloritic
blocks at its base.
Five hundred feet of rock containing tuff and lava flows then follow. At
the base lava rests with an irregular contact against cherty rocks at the top
of the underlying unit. This lava passes upwards into brecciated lava—
possibly a flow breccia then into a breccia more closely resembling the coarser
phases of the tuff. This is followed by another lava flow.
The remaining 460 feet of the formation commences with a very coarse
breccia (Plate x, fig. 4) which grades up into finer sediments. The next bed
is over two hundred feet thick, it has some fluctuations in grainsize but
becomes finer in the upper part. Above is an alternation of tuff and cherty
rock followed by seven feet of sediments with the bedding features of the
shaly rocks of the Waterbeach Formation. The formation concludes with a
hundred foot unit which grades from breccia at the base into cherty rocks
at the top.
Regional distribution and variation. South of the Turon River the
formation remains much the same as the type section, but to the north
there is lateral change in the lowest unit of the formation, where an increase
in the proportion of intercalated strata takes place and some of the tuffs
pass into conglomerates with cobbles of felsitic rocks. The lavas, too, in the
north differ somewhat in hand specimen from those in the type section;
vesicular types are common, the cavities being filled with epidote, prehnite
and calcite.
The general lithological identity and bedding characteristics of the
formation are maintained in the Limekilns area on the east side of the
Wiagdon Thrust where the formation is of the order of 1,500 feet thick.
There is a felsitic lava flow at the base to the north-east of Limekilns.
Overlying the tuff is a black shale (the Rosedale Shale) which is poorly
bedded and contains no tuffaceous material. The junction with the Merrions
tuff is sharp.
Petrography. Apart from minor differences, the sediments (Plate 111, fig. 1)
of this formation are the same as the tuffaceous rocks of the Turondale
Formation. Amphibole is still rare but is more common than in the Turondale
Formation. It has the following optical properties: Z = slightly greenish
mid-brown; X = very light brown; Z/\C about 17°; moderately large 2V.
Orthoclase, often partly sericitized, is common only in coarser tuffaceous
rocks; the mineral has more albite patches than in the type section of the
Turondale Formation. Comparison of the micrometric analyses of sediments
from the two formations (Tables 2, 4) and their positions on Figures 4 and 5
bring out this similarity. The mode of deposition of the tuffaceous rocks is
discussed above.
136 PALAEOZOIC STRATIGRAPHY AND SEDIMENTARY TECTONICS
The lavas associated with the Merrions Tuff are altered dacites; they
are tough dark grey to greenish grey rocks. The phenocrysts are mainly of
felspar (1:0 mm.), very rich in soda (about Aby;) and frequently corroded.
Quartz phenocrysts are not abundant but are always corroded. In one
thin-section a pale green non-pleochroic pyroxene was found. Amphibole
similar to that in the tuffs is occasionally present. The groundmass of
the lavas is composed of plagioclase laths (0-1 mm. long), granular quartz,
grains of epidote and wisps of chlorite. The degree of alteration in these
rocks is considerable; in most slides chlorite and epidote pseudomorphs
after amphiboles are common.
TABLE 4
Micrometric analyses of Tuffaceous Arenites from the Merrions Tuff
A B Cc D
Quartz .. he ae Be 3 27 10 8
Felspar.. i oe se 51 35 30 31
Igneous rock fragments : 6 - 19 3
Sedimentary rock fragments .. — — 5 ==
Chloritic patches 3 Re 4 7 = 3
Chloritic matrix ave 3 36 31 36 55
All specimens are from the type section of the formation.
> AtSalat 50 feet above the base of the formation.
5) AUS) 35 350 feet above the base of the formation.
TS 6. 850 feet above the base of the formation.
. TS 4. 1,030 feet above the base of the formation.
eholesi=
CUNNINGHAM FORMATION
This formation is named after the parish in which the type section occurs.
The formation consists of slates, siltstones, greywackes and conglomerates.
Lavas and tuffs are absent. Some of the greywacke beds are thicker than
those of the Waterbeach Formation but at the same time considerably
thinner than typical beds of the Merrions Tuff.
Type section. This is along the banks of the Turon River in Portion 138
of the Parish of Cunningham, conformably overlying the type section of the
Merrions Tuff. The stratigraphic column is given in Figure 9. The type
section which is a little over 2,800 feet has been measured to the top of
the highest bed exposed.
The proportion of sandy material and the grainsize of the coarser beds
is greater in this section, than in the type section of the Waterbeach
Formation. These two features have associated with them, an increase in
thickness of the coarsest beds and a less distinct alternation of sequences
of fine sediments with groups of greywacke beds than in the type sections
of the Waterbeach Formation and the middle part of the Turondale Formation.
In the Cunningham Formation type section, the coarser beds are more
abundant between 300 and 900 feet, 1,050 and 1,150 feet, 1,250 and 1,550 feet,
1,750 and 1,850 feet, and 2,130 and 2,850 feet above the base. The coarsest
greywackes in this formation are in thick beds—the thickest is 70 feet, one
is 60 feet thick and there are a number over 30 feet. This contrasts with the
type section of the Waterbeach Formation on which the thickest bed is 20 feet.
On the basis of number per hundred feet of section, thinner greywacke beds
are more abundant in the Cunningham Formation than in the Waterbeach
Formation. Most beds up to four or five feet thick are graded-bedded but the
thicker beds display irregular fluctuations of grainsize. The siltstones and
Slates are similar to those in the Waterbeach Formation.
G. H. PACKHAM 13
At two places in the type section, namely 1,450 and 2,150 feet above the
base, beds quite different in texture from the greywackes occur; the first
is 18 feet thick and the second is 50 feet in thickness. These are massive,
dark grey in colour, consisting dominantly of clay and silt size material
(see Table 5, Plate xt, fig. 2) but with a small proportion of coarser material.
LEGEND
Greywacke Slumped sediment
Interbedded silts & slates
1000’
300’
1TO0*
500‘ 15004 2500’
400° 1400’ 2400.
2300
1300’
ht
i!
lhl
hi
iN
\|
|
|
|
\!
\!
100‘ — 1100'+= 2100"
1000" 2000’
Figure 9. Type section of the Cunningham Formation measured on the south bank
of the Turon River, Portions 41, 50, Parish Waterbeach. Greywacke beds less than
2 feet thick are shown as thin horizontal lines.
In the lower one the largest pebbles are 2 to 3 inches in diameter, whilst
in the upper one the maximum size is 6 inches. The lower bed contains
deformed blocks of shale up to two feet in diameter in a matrix jwhich has
folded banding. One brachiopod fragment was found in this bed. The upper
bed is composed dominantly of a finer banded phase with about 80 percent
188 PALAEOZOIC STRATIGRAPHY AND SEDIMENTARY TECTONICS
clay and silt and a second phase which contains an appreciable proportion
of sand sized grains. The boundaries between the two phases are sharp but
very irregular (Plate x, fig. 6). Both are boulder bearing. The best explana-
tion of the origin of these two beds is that they are mudflow or highly mobile
slump deposits. The sporadic banding of the rock, the irregular distribution
of boulders through the bed, the incomplete mixing of the fine-grained and
sandy phases of the upper bed and the contortion of the banding of the
lower bed are all in agreement with this conclusion. The symmetry of the
contortions, incidentally, suggests that the direction of flow was from east
to west. The inclusion of the brachiopod in the lower bed suggests that the
flow was initiated in a fairly shallow-water environment.
The observations made regarding the origin of the alternation slate-rich
and greywacke-rich zones (p. 29) apply equally to the Cunningham Formation.
A voleanic origin can be definitely ruled out because there is no indication
of vulcanism during the deposition of the Cunningham Formation. It is
interesting to note that the upper mudflow horizon occurs at the base of a
greywacke cycle which is over 200 feet thick. This commences with thickly
bedded greywackes but the thickness of the beds gradually diminishes
upwards. This may have been caused by an uplift in the source area or fall
in sea level, erosion gradually obliterating the effects of the change and the
supply of sandy debris gradually diminishing.
Regional variation. The formation is confined to the western margin of
the area under discussion but some variation in the proportion of various
lithologies occurs. Greywackes with an abundance of shale fragments are
more common to the south and to the west there is a rapid decrease in the
abundance of coarse sediment.
Source direction of the sediments. Flute-casts and small-scale cross-
bedding occurring at intervals throughout the type section and the flow-
deformed banding in the lower mudflow horizon all indicate a direction of
sediment flow towards the west. Small-scale cross-bedding indicating a similar
direction of flow occurs near the base of the formation at Gimlet Creek (four
and a half miles due south of the type section) and again at several points
in the section exposed by Winburndale Rivulet.
Thus the same direction of slope of the sea-floor was maintained from
the time of deposition of the Cookman Formation until the deposition of
the Cunningham Formation.
Petrography and sedimentation. Beeause of the higher proportion of
shale fragments the appearance of the Cunningham Formation greywackes
differs from that of the Waterbeach Formation greywackes. The coarser
phases of the Cunningham greywackes may contain up to fifty percent of
shale fragments. The grey colour of the Cunningham Formation greywackes
contrasts with the greenish grey of those of the Crudine Group which contain
a higher proportion of epidote.
Micrometric analyses of some of the sediments of the Cunningham
Formation are given in Table 5 and are plotted in Figures 4, 5. The propor-
tion of felspar is significantly lower in the Cunningham Formation than in
the Waterbeach Formation. The thin-section characteristics of the various
types of detrital minerals and rock fragments are identical in both formations.
There are fragments of porphyritic acid igneous rocks, limestone, shale and
quartzite. A single pebble of granite found in a conglomerate phase of the
Cunningham Formation does not necessarily indicate any change in the
source area because a few granite pebbles have been found in the Waterbeach
Formation in the Hill Emde area. The siltstones and slates of the Cunningham
G. H. PACKHAM 139
Formation are poorly sorted; they differ from those of the Waterbeach
Formation only in that they contain a higher percentage of detrital quartz.
The percentage of matrix in the Cunningham greywackes is more variable
and often considerably lower than in the Waterbeach greywackes (Plate x1,
figs 4, 5), and the beds are often considerably thicker; only two percent of
the -greywacke beds in the type section of the Waterbeach Formation are
over 16 feet thick, while 10 percent of those of the Cunningham Formation
type section exceed this value. If the initial slope from the shallow-
water environment into the trough in which the Cunningham Forma-
tion was deposited, was steeper than for the Waterbeach Formation, and
if, the trough had a flatter floor, then I think the differences could be
explained. The steeper slope would allow flowing grain layers (Sanders,
1965) or turbidity currents with a smaller amount of fine material to flow;
the sediments found in the type section of the Cunningham Formation may
have been deposited at the foot of such a slope as a submarine fan. The
rarity of slumps in this section offers evidence against deposition on the
slope. Had the slope continued further to the west, it would be expected
that the coarse sediments would be common for a considerable distance in
this direction. This is not the case.
TABLE 5
Micrometric analyses of Greywarkes from the Cunningham Formation in the Sofala district
A B C -D E
Quartz ook AG ee 31 22, 10 20 2H
Felspar am ait sf ) 4 3 9 9
White mica aN sf — — 7 — —
Igneous rock fragments .. 25 32. 3 57 48
Sedimentary rock fragments 2 Sf 3 2
Matrix : Be ae 33 34 77 mt 14
A. TS .69. Type section, 330 feet above the base of the formation.
B. TS 61. Type section, 1,380 feet above the base of the formation.
C. TS 80. Type section, 2,150 feet above the base of the formation.
D. S 9. Sofala-Hill End road, $ mile west of the eastern bouadary of the formation.
E. $89. Sofala-Hill End road, ? mile west of the eastern boundary of the formation.
Fauna. Apart from a brachiopod fragment in one of the mudflow horizons,
no fossils have been found in this formation other than those occurring in
limestone pebbles in the coarser sediments. Tabulate corals are the most
abundant. The forms found are favositids, one is doubtfully Hmmonsia sp.,
the others are Favosites sp. One rugose coral, possibly an Acanthophyllid
has been found.
Age. The best evidence for the age of this formation comes from its
correlation with the Devonian Limekilns Group (see below). The Cunningham
Formation can be traced westwards to the Euchareena area where it passes
laterally into shallow-water sediments containing Lower Devonian fossils.
LIMEKILNS GROUP
This group consists of three formations which overlie the Merrions Tuff
in the vicinity of Limekilns, five miles south-east of Wattle Flat. These three
formations were first described by Hawkins (1953). In ascending order, the
formations are the Rosedale Shale, the Jesse Limestone and the Limekiln
Creek Shale. There is no indication of contemporaneous vuleanism within
the group though it is overlain and underlain by thick pyroclastic formations,
the Winburn Tuff and the Merrions Tuff respectively.
140 PALABOZOIC STRATIGRAPHY AND SEDIMENTARY TECTONICS
The age of the limestone in this group has been known to be Devonian
for many years but the limestone was not mapped stratigraphically until it
was studied by Hawkins (1953). The Limekilns Group has an important
place in the interpretation of the palaeogeography of the region since it
contains neritic benthonic organisms in their living environment and is of
the same age as the Cunningham Formation which is a typical geosynclinal
deep-water succession. The evidence for this correlation rests on a number
of different lines of approach. First, they both overlie the Merrions Tuff
conformably. Then the lithologies are comparable, in that they are both non-
voleanic; this is not a common feature in the succession. Again, sedimentary
structures in the Cunningham Formation in the Turon River section show
that the sediment has been transported from east to west, indicating the
existence of an area with a shallow water environment somewhere to the east.
The thickness of the Limekilns Group in the Limekilns area is of the
order of 2,500 feet. This may be an overestimate because faults are difficult
to detect in the Rosedale Shale and the Limekiln Creek Shale.
a. Rosedale Shale. This formation which has been named after a property
at Limekilns, has its type section (as. designated by Hawkins, 1953) in
Pender’s Creek in Portions 1, 16 and 63 of the Parish of Jesse. A sharp
contact with the Merrions Tuff marks the lower boundary. The shale is
dark grey to black, well jointed and fisile but with macroscopic bedding
planes every six to eight inches. Pyrite is common. The shale is composed
of quartz and felspar silt set in a fine-grained matrix of chlorite and sericite.
Chlorite is the more abundant. Fossils are rare, there are only occasional
linguloid brachiopods and poorly preserved plant remains. The thickness
quoted by Hawkins for this formation is 700 feet.
b. Jesse Limestone. This formation conformably overlies the Rosedale
Shale. The outcrops to the east of Limekilns are thickly bedded limestones and
breccias, while those to the west are dominantly calcarenites 50-100 feet thick,
in which individual beds can be traced for distances of at least a mile. Hawkins
(1953) designated the section of the clastic facies exposed in Diamond Creek
as the type section. The base and top of the formation in the type section
are respectively the base of the first calcarenite bed and the top of the
highest one.
The limestone is highly fossiliferous, though the massive limestone is
less abundantly so than the detrital facies to the west. Syringaxon ? sp.,
Phillipsastraea currani, Hexagonaria tunkanlingensis, Disphyllum sp., Dendro-
stella sp., Spongophyllum sp., Acanthophyllum mansefieldensis, Grypophyllum
sp., Lyrielasma sp., Tryplasma sp., Pseudampleaus sp., Plasmophyllum
australe, Calceola sp. and Receptaculites australis have been identified by
Wright (1966). Brachiopods, stromatoporoids, tabulate corals, fish plates
and conodonts are also known from the formation. Preliminary identifications
of the conodont fauna are: Bellodella devonica, B. resima, Hindeodella
prescilla, Lonchidina sp., Neoprioniodus excavatus, Neoprioniodus sp., Ozar-
kodina denckmanni, Paltodus acostatus, P. unicostatus, P. valgus, Polygnathus
linguiformis foveolata, Spathognathodus exiguus, Synprioniodina sp. and
Trichonodella sp. The assemblage is apparently a late Lower Devonian one
(Emsian).
Towards the north, along the Limekilns-Wattle Flat road, the limestone
thins very considerably and the formation is represented by a few impure
calcareous arenites. These contain, apart from detrital carbonate, porphyritic
acid igneous rock fragments as the most abundant constituent with quartz
grains less abundant and felspar least abundant. This the same order of
“I G. H. PACKHAM 141
abundance as the various types of detritus in the greywackes of the
Cunningham Formation.
ce. Limekiln Creek Shale. This formation was also defined by Hawkins
(1953). It has its type section in Cheshire’s Creek in Portion 253 and the
eastern part of Portion 127 of the Parish of Wiagdon. The thickness of
the formation given by Hawkins is 1,500 feet. It overlies the Diamond Creek
Limestone and underlies the Winburn Tuff. The lithology and fossil content
are similar to those of the Rosedale Shale.
WINBURN TUFF
The formation has its type section in Cheshire’s Creek, in Portion 127
of the Parish of Wiagdon (Hawkins, 1953). The formation is named after
a parish which has Cheshire’s Creek as its boundary and so the south bank
of the creek in the type section is actually south of Limekilns, in the Parish
of Winburn. The dips of the underlying shales are frequently discordant
with the boundary of the tuff which may lie unconformably on the Limekilns
Group.
The formation occurs in the core of a synclinal structure truncated to
the west by a thrust fault. The section is thus incomplete but over 2,000 feet
of it is preserved. Lithologically, the formation is very similar to the Merrions
Tuff. The beds are thick and the greatest part of the formation is composed
of tuff of coarse sand size. The remainder is composed of finer tuffaceous
sediments.
No indication of vulcanism has been found within the Cunningham
Formation and this is taken as an indication that the Cunningham Formation
pre-dates the Winburn Tuff and so may be equated with the Limekilns Group.
There is no formation to the west with which the Winburn Tuff can be
correlated.
There is a general textural and mineralogical resemblance between the
Winburn Tuff and the Merrions Tuff. The rocks are poorly sorted and com-
posed of white and pink. felspar, quartz, epidote and chlorite; rock fragments
are not abundant. Orthoclase is the dominant felspar, it is often partly
replaced by albite in patches about a tenth of a millimetre long or less
commonly, by narrow veins along the cleavage. The detrital plagioclase in
the rock is albite. Epidote is less abundant than in the Merrions Tuff.
THe STRATIGRAPHY OF THE Hitt ENnp District
There have been three previous geological maps of the Hill End region,
each covering an area of about 100 square miles surrounding the town. The
first was by Pittman (1881); his map showed that the town, and thus the
mineralized zone, lay on the crest of an anticline, but he made no attempt
to subdivide the succession. The second map was produced by Harper (1918)
who, because he misinterpreted some sedimentary structures in the succession,
considered that an unconformity was present. The third study was by Jopling
(1950) who reaffirmed the original simple concept of Pittman, but again there
was no serious attempt to divide the sequence into units, since the purpose
of Jopling’s study was to investigate the gold mineralization. The anticline
on which the town of Hill End is situated gives the most complete section
in the central part of the region. The sequence of formations from the
Chesleigh Formation to the Cunningham Formation, described in the Sofala
district can also be recognized in the Hill End area. The geological map
accompanying (Plate 1v), incorporates the Hill End area.
142 PALAEOZOIC STRATIGRAPHY AND SEDIMENTARY TECTONICS
CHESLEIGH FORMATION
The lowest part of the section consists of massive subgreywackes, with
tuffs and thin beds of slates at the base. Above this, massive tuffaceous rocks
appear which have sorting akin to that of greywackes; these tuffaceous rocks
are reminiscent of those in the Turondale Formation of the Crudine Group,
but their association with massive subgreywackes is one unknown in that
formation to the east. The exposed thickness of the formation in the Hill End
area is about 1,700 feet, in all probability representing the upper part of the
Chesleigh Formation which, in the type area, contains volcanic material.
Unfortunately, it has not been possible to obtain any evidence of the
direction of transportation of the sediments in the Chesleigh Formation out-
cropping in the Hill End area; the direction of source of the Chesleigh
Formation in the Sofala-Wattle Flat area is from the west or a little to
the south of west. The Hill End area would be expected to be a little closer
to the source area; this seems to be reflected in the lithology of the sediments.
The non-voleanic sediments are more felspathic; a notable addition is the
presence of a small amount of microcline. There is also a suggestion of
coarsening of grainsize, indicated by a greater abundance of shale blocks
in the subgreywackes. Volcanic material is also more abundant here than
in the type section, again suggesting a closer proximity to the source region.
The outcrop in the Hill End district, is confined to the axial region of
the Hill End Anticline, and occupies a strip, less than two miles in width,
commencing two miles north of the town of Hill End and extending south
for 16 miles. The outcrop is cut off to the north and the south by the gentle
plunge of the fold.
CookMAN FoRMATION
In the Hill end Anticline, the Chesleigh Formation is flanked by the
Cookman Formation with an outcrop width of just under half a mile on
both limbs. Good cliff sections exist where the Cookman Formation crosses
the Turon and Macquarie Rivers. The Cookman Formation also outcrops
to the east of the Hill End Anticline in the axial regions of three small anti-
clines along the Macquarie River upstream from its junction with the
Winburndale Rivulet. Only the upper part of the formation is exposed in
each case. The thickness of the Cookman Formation in the Hill End area
is about 2,300 feet, i.e., considerably thicker than in the type section (1,500
feet). The coarser sediments like those of the type section, are almost all
quartz-rich with a few percent of argillaceous matrix; they occur in beds
mostly one to two feet thick. Unlike the type section, graded-bedding is well
developed although the size range is rather more restricted here. These
features suggest a location for the Hill End sections of the Cookman Forma-
tion further from the source of the sediments. The indications in the Sofala
area were that the sea-floor sloped to the west during deposition of the
Cookman Formation in contrast with the general easterly slope at the time
of deposition of the Chesleigh Formation. In the Turon River cliff section
of the formation exposed on the eastern side of the Hill End Anticline
slipped load-casts, small-scale cross-bedding and slumps are occasionally
found and these indicate a westerly slope on the seafloor.
In hand specimen, the subgreywackes of this formation differ in appear-
ance from those in the Sofala area. This difference results from the greater
degree of metamorphism in the Hill End area. Thus, instead of the rock
being moderately indurated and quartzitelike with individual grains fairly
distinct, it has been converted to a very hard, brittle, somewhat translucent
and hornfels-like rock in which the grains are entirely interlocking. Under
the microscope it is difficult to determine the nature of the original texture.
st G. H. PACKHAM 143
CRUDINE GROUP
Both the Turondale and the Waterbeach Formation can be traced into
this area and maintain many of the characteristics found in their type
sections. .
Turondale Formation
This overlies the Cookman Formation in the Hill End Anticline and out-
crops over the whole length of the anticline in the area mapped. Although
it does not appear in any of the anticlines to the west of the Hill End
Anticline it does outcrop extensively east of it, in the anticlinal structure
west of the junction of the Macquarie River and the Winburndale Rivulet.
This outcrop has a width of three miles on the southern border of the map,
interrupted only by three small inliers of the Cookman Formation. To the
north this anticlinal structure plunges north so that the Turondale Formation
disappears under higher formations.
The thickness of the formation seems to be fairly constant in the region
and of the same order as the thickness measured in the Turon River section
at Hill End, that is, 2,200 feet (in the type section it is 2,000 feet). The
lithology likewise is similar—mainly tuffaceous rocks and greywackes. The
beds are thick but for the most part the coarse rocks characteristic of the
lowest part of the formation in the type section are not present. This means
that the junction of the Turondale Formation with Cookman Formation is
not such a distinct one as in the Sofala area but it can be recognized by
an increase in the thickness of beds, the presence of some beds of coarse
sand size and the presence of acid tuffs and most important, the change from
dominantly quartz detritus to lithic and felspathic debris. Although there
has been a reduction of the maximum grainsize of the sediments in this
‘formation in the Hill End area, the abundance of coarse-grained sediments
is far greater here than in the Waterbeach Formation. Some of the thicker
beds of the Turondale Formation contain blocks of shale—one of these
observed in Washing Gully of Hill End is ten feet long. Indications of
slumping movements are often seen in these beds. The tuffaceous rocks
range from coarse sand-size through to fine cherty types.
A porphyry, twenty to thirty chains wide in outcrop, occurs within the
Turondale Formation, at the same stratigraphic level, on both sides of the
Hili End Anticline. Even where its contacts are exposed there is no clear
evidence that the porphyry is intrusive and, further, it has obviously been
involved in the folding movements since it is strongly deformed. It may be
a flow. An inclusion of granite with a rounded outline, measuring approxi-
mately two feet by one foot is exposed in the porphyry in the bed of Washing
Gully a mile and a quarter west of Hill End. The inclusion shows some
signs of disruption for small groups of crystals, evidently derived from the
granite, are seen separated from the inclusion by a narrow vein .of the
porphyry. There is no indication of the development of any kind of reaction
rim around the inclusion.
The evidence of the direction of slope of the sea-floor is in harmony with
what has been found in the Sofala area. The general indication is that the
slope was from east to west, which means that the Hill End section was
further from the shore. This accords with the lithological data such as the
absence of the coarsest tuffs and the increase in proportion of greywacke.
The data indicating the slope are a slump structure in tuffaceous sediments
on the Macquarie River three miles south-east of the junction of the Turon
and Macquarie Rivers and the occurrence of cross-bedded ripple marks near
144 PALAHOZOIC STRATIGRAPHY AND SEDIMENTARY TECTONICS
the base of the formation 300 yards west of the same river junction. Small-
scale cross-bedding exposed on the bank of the Macquarie River, 15 miles
south of Hill End on the western limb of the Hill End Anticline, indicates
an easterly downward slope of the sea-floor. Thus, the axis of the trough in
which the Turondale Formation was deposited lay in the vicinity of the Hill
End Anticline.
Waterbeach Formation
This formation outcrops extensively in the Hill End area. It appears
on both limbs of the Hill End Anticline and, on the eastern side towards the
south of the area, can be traced around several folds to the junction of
Winburndale Rivulet and the Macquarie River. About 6 miles west of the
Hill End Anticline the Waterbeach Formation outcrops in the axial region
of the Ulmarrah Anticline (Figure 1).
The appearance of the Waterbeach Formation in most sections is closely
similar to that in the type section with the notable exception of the inher
in the Ulmarrah Anticline described below. The usual features are: an
absence of tuffs, a preponderance of slates over greywackes, and well developed
graded-bedding, especially in the slates. This graded-bedding is better
developed than in the type section. The same tendency characterizes the
Cunningham Formation in this area.
In a number of places in the section of the Waterbeach Formation
exposed on the western limb of the Hill End Anticline, flute casts and small-
scale cross-bedding indicate a down-slope of the sea-floor from east to west.
Hence, the same direction of slope persisted from Sofala at least as far west
as this section.
The thickness of the Waterbeach Formation south of Hill End on the
Turon River is 2,000 feet, 350 feet thicker than the type section.
In the Ulmarrah Anticline immediately underlying the Merrions Tuff
there is over 900 but less than 1,500 feet of extremely deformed sediment.
This material is part of a huge slump, the base of which is not exposed.
The sediment in the deformed zone is coarse greywacke and slate, which have
been mutually involved in the slip. The greywacke (Plate x1, fig. 7) is not
a normal one; it contains a proportion of matrix much larger than is normal
and the sorting is poorer (cf. Table 6). In the field it is also unusual, for
included in it are blocks of a great variety of rocks, the largest and
commonest being shale. Blocks 15 feet across are common. Other inclusions
are not only far less frequent, but of considerably smaller size, being for the
most part rounded boulders up to 2 feet in diameter. Rock types represented
are: quartz-felspar porphyries, biotite-granite, limestone, dacitic tuff,
quartzite and quartz. In places the boulder-bearing material and deformed
Slates are overlain by little-deformed graded-bedded slates but elsewhere the
Slate is clearly injected by irregular masses of the greywacke (Plate x, fig 7) ;
these injection movements clearly pre-date the regional cleavage. These
relationships may be the result of first, the slumping of the coarse material
into a position of temporary stability, then deposition of graded-bedded shales
on top followed by a later slip of both the first slump and its cover. This
process may have been repeated a number of times. Some of the contact
between the two phases might represent a slide-plane within the whole slump
mass. As it has not been possible to recognize any systematic arrangement
of folds within the slumped material, no direct evidence of the direction of
source of the slump is available.
G. H. PACKHAM 145
By far the best exposures of the slump mass are in the gorge of Pyramul
Creek on the northern edge of the area mapped. To the south, outcrops in
the axial region of the anticline are poor but the occurrence of poorly sorted
ereywackes and boulder beds, suggests that the slump extends for at least
six miles along the axis of the anticline. Further south the Waterbeach
Formation is concealed by the overlying Merrions Tuff. To the east, in the
Hill End Anticline, there is no certain indication of the slump, despite good
exposures. The only possible disturbed area is on the west limb of the anti-
cline on Pyramul Creek where some of the interbedded slates and greywackes
are rather more folded than usual; this folding, however, might be tectonic.
Although the evidence for the direction of origin of the slump mass is negative,
it suggests that the slump was derived from the west since, if it had come
from the east the effect certainly would be clearly noticeable in the exposures
in the Hill End Anticline. Again, if the slump had moved from the east
to the west the distance of movement would have to be very considerable
since boulder-bearing horizons are rare in the Waterbeach Formation within
the area mapped. Since sediments in the Waterbeach Formation, on the
TABLE 6
Micrometric analysis of Greywacke Matrix of
Ulmarrah Slump
Pt
Quartz .. tes a ae 5 22
Felspar ot at a oe 12-7
White mica .. ae il ys 2°8
Rock fragments : Bee es 13-8
Matrix se ait re ae 58-2
Locality : Axis of Ulmarrah Anticline in Pyramul
Creek.
western limb of the Hill End Anticline, indicate a westerly slope of the sea-
floor, the axis of the trough in which the formation was deposited must have
lain in the vicinity of the Ulmarrah Anticline or slightly to the east of it.
Another boulder-bearing greywacke, this time a very thin bed, by
comparison with the slump described above (the thickness of sediment
involved is only of the order of tens of feet), outcrops in the core of an
anticline some twelve miles south-east of the Ulmarrah Slump but is east of
the Hill End Anticline and so is unrelated to the large slump.
MERRIONS TUFF
In the Hill End Area, this formation maintains the character of its type
area at Sofala better than the underlying units. North of the Turon River
it outcrops on both the eastern and western limbs of the Hill End Anticline
but to the south occurs only along the western limb. On the eastern limb
the Merrions Tuff outcrops over a considerable area between the Maequarie
and Turon Rivers due to the northerly plunge of several folds of moderate
amplitude west of the Hill End Anticline. East of these structures the
formation again strikes southwards along the western side of the synclinal
structure continuing south from Sally’s Flat. To the west of the Hill End
Anticline the tuff outcrops on both limbs of the Ulmarrah Anticline as far
south as the Macquarie River. The unit again occurs in a southerly-plunging
anticline on the northern edge of the area mapped, between the Hill End
Anticline and the Ulmarrah Anticline. The Merrions Tuff in the Ulmarrah
Anticline is the most westerly outcrop of the formation in the region under
discussion. The area east of the Hill End Anticline is structurally different
J
146 PALABOZOIC STRATIGRAPHY AND SEDIMENTARY TECTONICS
from other areas in which the tuff outcrops for here it is tightly folded into
sharp crests and troughs whereas elsewhere it behaves in a more competent
fashion.
Lithological and bedding characteristics similar to those of the formation
in the type section are maintained in the Hill End area, where the thickness
is over 2,000 feet on the eastern limb of the Hill End Anticline. To the west
of Hill End the formation contains less coarse material and the section is
thinner (about 1,000 feet at Ulmarrah). Altered dacitic lavas are present
in nearly all exposed sections and generally they occur at, or near, the base
of the formation. The lavas sometimes have columnar jointing and, as a
result of the regional deformation in which they have been involved, the north-
south dimension of the columns is generally one and a half times the east-west
dimension.
The lithological continuity of the Merrions Tuff and the decrease in
thickness to the west are such that it must have been derived from the east.
A shift to the west of the axis of the trough must be postulated to account
for the extension of the sediment as far west as the western limb of the
Ulmarrah Anticline. During the time of deposition of the underlying Water-
beach Formation the axis lay in the vicinity of the Ulmarrah Anticline.
CUNNINGHAM FORMATION
The type section of the Cunningham Formation which is described above
is on the western side of a synclinal structure running through Sally’s Flat.
South of the Turon River the formation is restricted by the complex anticlinal
structure occurring between the river and the Hill End Anticline but north-
wards the formation extends across to the anticline. West of the Hill End
Anticline the Cunningham: Formation outcrops over a very large area in a
broad synclinal structure (the Ophir Syncline, Figtire 1) roughly ten miles
wide. The continuous outcrop is broken only by the lower rocks in the
Ulmarrah Anticline and the small structure between it and the Hill End
Anticline. The Cunningham Formation can be traced west to Euchareena;
the outcrops there are discussed below.
The thickness of the Cunningham Formation in the Ophir Syncline is
of the order of 12,000 feet. This is far greater than the thickness of the type
section (2,800 feet). But it is not certain whether the formation has thickened
to the west since there is no formation ovenl yang the type section of the
Cunningham Formation.
The lithology of the Cunningham Formation in the Hill End area differs
from that of the type section principally in the proportions of the various
rock types. Fine-grained sediments are far more common here and conglo-
meratic greywackes are very rare. To the west of Hill End greywackes are
rare except near the base of the formation and there they are rich in felspar
apparently derived from tuffaceous rocks. Between 5,000 and 6,000 feet above
the base of the formation, a few thin graded-bedded and slumped bands of
arenite and rudite composed largely of calcareous debris, outcrop on the banks
of the Macquarie River between Curragurra Creek and Pyramul Creek. The
detritus was apparently derived from the west where the Cunningham
Formation passes into limy sediments. The formation here consists almost
entirely of slate. Individual bands of silt are not common. Almost all of the
silty material occurs at the base of graded slaty beds. These graded units
are thicker than those described as occuring in the type section of the Water-
beach Formation. The general order of thickness of -beds is six inches in the
Cunningham Formation in this area.
a) G. H. PACKHAM 147
The location of the axis of the trough in which the formation was
deposited seems to have been in the vicinity of the Hill End Anticline. This
is suggested by two lines of evidence. First, the shale-pebble conglomerates
which are so characteristic of the formation in the eastern part of the region
only extend as far west as the eastern limb of the Hill End Anticline.
Secondly, there are frequent slump structures west of the Hill End Anticline ;
these make their appearance on the western limb of the anticline and are
found commonly in most sections to the west of it. Good examples of these
slump structures are exposed in Tambaroora Creek upstream from Washing
Gully and at a number of places along the Macquarie River between Tamba-
roora Creek and Lewis Ponds Creek west of the axis of the Hill End Anticline.
No direct evidence of the direction of movement of the sediment has been
obtained by an examination of the structures because the amount of move-
ment has almost completely destroyed the bedding. The deformed material,
largely of clay and silt size, is often cut by veins of silty material probably
deposited while water was being squeezed from the plastic mass after its
final deposition. The indications of graded-bedding and other structures,
which remain suggest that the initial deposition of the sediment was in deep
water. The fine-grained nature of the sediments involved in these slumps and
the absence of similar structures on the eastern side of the Hill End Anticline
suggest an easterly slope of the sea-floor as far as the Hill End Anticline.
Convolute bedding is common locally in very fine silts just west of the
Ulmarrah Anticline and near the junction of Lewis Ponds Creek and the
Macquarie River.
THE STRATIGRAPHY OF THE EUCHAREENA DISTRICT
The Devonian limestones and some of their associated sediments in the
valleys of Nubrigyn and Boduldura Creeks, west of Euchareena and Stuart
Town have been mapped and petrographically examined by Wolf (1965).
Apart from this, there have been only two significant contributions to the
geology of the area, the original mapping by Carne and Jones (1919) of the
limestones studied by Wolf and the reconnaissance mapping of the western
margin of the area by Joplin and others (1952).
OAKDALE FORMATION
The oldest rocks in the Euchareena district are andesitic volcanics,
representing a southern extension of the Oakdale Formation from the Mumbil
District (Strusz, 1960). The Ordovician age of the volcanics has been estab-
lished by the finding of graptolites at four localities. The northernmost is
in Portion 10, Parish of Nubrigyn. Sherrard (1954) states that the graptolites
here are: Climacograptus bicornis, Climacograptus scharenbergi, Ortho-
graptus cf. apiculatus and Lasiograptus ? harknessi. Two miles to the south
on the boundary of Portions 4 and 48 of the same parish there are fragments
of Dicellograptus sp. Further south again on the Bell River just east of the
Silurian Nandillyan Limestone (Joplin and others, 1952), I have found
Orthograptus cf. wpiculatus and Glossograptus hinksti. The fourth locality
is 21 miles west of Mullion Creek Railway Station, by the side of the
Belgravia Road. The forms identified are Orthograptus apiculatus and
Climacograptus sp.
These andesitic volcanics occupy the same position in the succession as
the Sofala Volcanics and are of comparable age. There is some difference in
the general appearance of the rocks in the two areas. The andesitic volcanics
in the western area are far less indurated than the Sofala volcanics and
the interbedded fine-grained rocks are shales rather than chert.
148 PALAEOZOIC STRATIGRAPHY AND SEDIMENTARY TECTONICS
An important section occurs on the eastern margin of this formation in
the northern-western corner of the area mapped, in a gully flowing into
Nubrigyn Creek through Portion 10 of the Parish of Nubrigyn. The section
is as follows:
The lowest rocks exposed are andesites, overlying these is a thin limestone
followed by shales with the fauna containing Climacograptus bicornis men-
tioned above; this in turn is followed by marls and a second thin limestone
containing a fauna of tabulate corals overlain by a thick succession of arenites
consisting of material derived from andesites and finally a Silurian lime-
stone at the base of the Mumbil Formation, containing Pentamerids,
Tryplasma sp., and a rugose coral close to Phaulactis shearsbyi. The exposed
section below the Mumbil Formation is about a thousand feet thick.
The fauna of the lowest limestone in this sequence is highly significant ;
it includes halysitid corals, Syringopora sp., Heliolites sp., and Multisolenia
sp. A comparable fauna has been found in the Oakdale Formation, about
thirteen miles to the north (Strusz, 1960). The graptolite assemblage has
been placed by Sherrard (1954) in the zone she calls the Zone of Climaco-
graptus peltifer but there is no clear reason why it should be placed in that
zone. The fauna is compatible with the assemblage two zones higher in what
she calls the Zone of Orthograptus calcaratus and Plegmatograptus nebula.
Thus the field observations indicate that the Halysites fauna extends down
into the Upper Ordovician at least as far as the last-mentioned zone.
MULLIONS RANGE VOLCANICS
In the Mullions Range this formation occupies the axial region of an
anticline plunging to the north. Unfortunately no older formations outcrop
in the structure so that it has not been possible to determine the thickness
of the formation in the Mullions Range. The formation outcrops to the west
of the Mullions Range as a gradually narrowing strip, running slightly west of
north. It has been traced as far north as Eadvale.
There are at least 5,000 feet of the volcanics exposed in the Mullions
Range, but in the wwell-exposed section along Kerr’s Creek, chosen as the type
section, the volcanics are only 1,500 feet thick. The thickness of the formation
diminishes rapidly towards the north. In the type section there is a dacite
flow at the base, followed by breccias of similar composition and then another
dacite flow. On top of this are more clastic sediments—tuffs and coarse
sandstones. The section ends with a flow of banded rhyolite. The changes
in thickness within the formation seem to be an original feature, since there
is no local indication of an erosional break at the top. A volcanic centre
in the vicinity of the Mullions Range, may account for the distribution of
the formation. The Mullions Range Volcanics rest on the Ordovician andesitic
rocks described above (Oakdale Formation).
The volcanics are almost exclusively dacites and rhyolites. The coarsest
rocks are porphyry-like types; these grade in grain-size to types containing
only a very small percentage of minute phenocrysts. The fine-grained types
are difficult to identify positively in the field because of their similarity to
the inter-bedded, indurated clastic rocks. The fine-grained lavas are pale
greenish-grey when fresh and only very rarely banded, so very few dips and
strikes can be obtained within the outcrop of the formation. The phenocrysts
in the lavas are orthoclase, plagioclase (albite) and quartz. Felspar and
quartz are roughly equal in abundance. The groundmass is composed of
interlocking grains of quartz and felspar, with only minor amounts of biotite
(X = pale yellow-brown and Z= very dark brown), mostly altered to chlorite.
Epidote is common, occurring as grains in the groundmass.
a G. H. PACKHAM 149
The clastic sediments in the formation are all highly indurated. They
range from fine silts to coarse sandstones and are composed of material
derived from the lavas. Glass shards are common in some of the tufts.
The age of this formation will be discussed after the description of the
overlying formations.
Mumesit ForRMATION
The Mumbil Formation originally described by Strusz (1960) in the
Mumbil district, conformably overlies the Mullions Range Volcanics. It has
been mapped from the northern extremity of the area, south to Kerr’s Creek,
around the margin of the Mullions Range in the Parish of Trudgett and then
down the eastern side of the range to Frederick’s Valley Creek. In the
northern part of the area the formation rests directly on rocks older than
the Mullions Range Voleanics.
The section in Frederick’s Valley Creek is 1,200 feet thick consisting
of grey and greenish-grey slate, commonly containing large pyrite cubes, and
is almost devoid of bedding. The cleavage developed in the type section is
not typical of the formation as a whole. To the west, the cleavage gradually
diminishes in intensity, from Nubrigyn Creek to Kerr’s Creek it is virtually
absent. The formation is non-volcanic and composed of fine-grained sediments.
The base of the formation, where it rests on the Mullions Range Volcanics
is taken as the top of the last lava or tuff band of the volcanics. At the
northern extremity of the area the formation rests on the Oakdale Formation.
The upper boundary is the base of the first tuff band of the Bay Formation
in the vicinity of the Mullions Range. In the Nubrigyn Creek area where the
thickness is only about 300 feet, the base is the lowest tuff band or andesite
flow of the Cuga Burga Voleanics. Just to the west of Kerr’s Creek where
neither the Mullions Range Volcanics nor the Cuga Burga Voleanics is
present, the mapping of the Mumbil Formation is very difficult since it is
overlain by the Cunningham Formation which contains few sandy bands here.
The top of the Mumbil Formation has been taken in this area as the place
in the sequence where the greenish colour of the claystones gives way to the
grey shales and siltstone bands of the Cunningham Formation.
A number of limestone lenses have been recognized within the Mumbil
Formation as shown on the map; they are probably equivalent to the Narragal
Limestone (Strusz, 1960). The fauna of the northern-most one includes
Tryplasma sp., Phaulactis shearsbyi and pentamerid brachiopods.
Bay ForMATION
The Bay Formation outcrops around the northern and eastern flanks of
the Mullions Range. Outcrop commences near Kerr’s Creek Railway Station
passing around the northern end of the Mullions Range, along the eastern
side through Bay Trigonometrical Station (after which the formation is
named), and then south to Frederick’s Valley Creek. The Bay Formation
overlies the Mumbil Formation and underlies the Cunningham Formation
but there is apparently an erosional break between the Bay Formation and
the overlying Cunningham Formation. West of Kerr’s Creek the Bay
Formation is absent and the Cunningham Formation rests on the Mumbil
Formation. The type section of the Bay Formation is exposed in Curragurra
Creek at the northern extremity of the outcrop of the formation. The thickness
in this section is approximately 400 feet, from here the thickness increases
steadily along the eastern margin of the Mullions Range reaching about 1,500
feet at Frederick’s Valley Creek. The proportions of the various lithologies
making up the formation change throughout its outcrop. The type section
150 PALAEOZOIC STRATIGRAPHY AND SEDIMENTARY TECTONICS
is representative of the northern and more westerly part of the formation.
It consists dominantly of tuffs of dacitic composition lithologically very
similar to the variety of types found in the Merrions Tuff, ranging from
coarse-grained crystal tuffs with chlorite-rich patches, to fine-grained chert-
like types with a conchoidal fracture. A small proportion of siltstones and
slates are present. By contrast, sections exposed to the south-east contain
a higher proportion of slates and silts. In Frederick’s Valley Creek, tuffs
are less abundant than slates and silts and some greywackes are present.
The top of the formation is taken as the top of the last tuff bed. It is overlain
by greywackes, conglomerates and slates of the Cunningham Formation.
Cuca Burca VoLcANIcs
Like the Bay Formation, the Cuga Burga Volcanics conformably overlie
the Mumbil Formation. The volcanics occur only in the north-western corner
of the area, extending south as far as the head of Weandre Creek. The
volcanics have been traced intermittently into the Mumbil District whence
they were first described (Strusz, 1960). The best exposed section in the
Euchareena district is seen in the vicinity of Eadvale, in the east-west gully
in Portion 58 of the Parish of Nubrigyn. The exposure there consists of
about 1,000 feet of andesite flows which exhibit pillow structures in places,
overlain by about 500 feet of interbedded shales and tuffaceous rocks. These
latter are similar to the tuffs of the Bay Formation, containing abundant
felspar, some quartz and chlorite. The beds of tuff are thin, of the order of
one to three feet thick. The top of the formation is the uppermost bed of tuff.
CUNNINGHAM AND “NUBRIGYN” FORMATIONS
The Cunningham Formation is by far the most extensive stratigraphic
unit in the Euchareena area, outcropping from Nubrigyn Creek to the
Macquarie River. Limestones, impure calecarenites and conglomerates inter-
digitate with the western margin of the outcrop of the Cunningham
Formation. The limestones were mapped by Carne and Jones (1919) who
gave them the name of the Nubrigyn (limestone) belt. All the limestone
lenses were collectively called the Nubrigyn Limestone by Packham (1958).
Wolf (1965) modified this term by including the clastic calcareous and lithic
sediments as well as the limestones in the Nubrigyn Formation. The southern
and eastern limits of the formation have not been defined closely, so at
present it seems best to use the term “Nubrigyn Formation” provisionally
until its stratigraphic and geographic limits are established. The approximate
limits of the rocks of “Nubrigyn Formation” lithology are shown on the
regional geological map.
The basal beds of the Cunningham-“Nubrigyn” sequence along the western
margin of the outcrop are shales which are overlain by interbedded calcareous
labile sandstones, polymictic conglomerates and shales, containing limestone
lenses which Wolf (1965) regards as algal bioherms. The limestone lenses
occur abundantly over about six square miles mainly south of Boduldura
Creek but extend south over about nine miles. Wolf (1965) divided the
Sequence above the basal shale into four units in the Boduldura Creek area.
The lowest consists of well-bedded impure calcareous sandstones, calcarenite,
shale and large algal bioherms. The second is not so well bedded, the algal
bioherms are smaller and more frequent, local areas of andesite occur (? flow
remnants) and there are limestone breccias developed around some of the
limestone pods. The third unit contains abundant voleanic detritus and
further ? flow remnants. Current bedding is common in the arenites. The
highest part of the exposed section is an algal bioherm. The thickness of
these strata preserved in this area is of the order of 1,700 feet.
rf G. H. PACKHAM 151
Much of the calcareous material in the “Nubrigyn Formation” contains
obscure organic structures which Wolf (1965) regards as algal. From the
better preserved material Johnson (1964) has described the following algae:
Hedstroemia australe, Garwoodia primitiva, Litanaa robusta, L. cracens,
Abacella deliculata, Lancicula wolfi, Uva sp., Litopora spatiosa, Girvanella
sp. aff., Girvanella ducti, Rothpletzella devonicwm and Renalcis devonicus.
Johnson also records the presence of the encrusting foram Wetherdella. Many
other fossil groups are present, tabulate and rugose corals are common,
stromatoporoids, brachiopods and conodonts have been found. The coral fauna
recorded by Strusz (1968) is Acanthophyllum (Neostringophyllum) impli-
catum, Calceola sp., Eridophyllum immersum, Heaxagonaria approximans
cribellum, Pseudochonophyllum pseudohelianthoides and Xystriphyllum
dunstani. Pseudamplexus princeps, Tryplasma spp. and Receptaculites sp.
are also present. A number of conodonts have been isolated from beds in
the lower part of the formation 0-6 miles north-north west of the junction
of Boduldura and Nubrigyn Creeks. The forms present include Spatho-
gnathodus linearis, S. imclinatus wurmi, S. cf. steinhornensis, Icriodus
pesavus, Ozarkodina sp. cf. O. jaegeri, O. media, O. denckmanni, Trichono-
della sp., Neoprioniodus sp., Hindeodella sp. and Panderodus wnicostatus.
The age of the formation is Lower Devonian, its precise position is discussed
later in this paper.
The other material in these sediments is igneous and metamorphic;
fragments of andesites, basalts, dolerite, fine-grained acid volcanic rocks,
quartzite and granite are all present. The first two were probably derived
from the underlying Oakdale Formation, the Cuga Burga Volecanics or
possible contemporaneous flows. The dolerite fragments are very altered and
very little can be determined of their original petrography. Dolerites are
known to intrude Ordovician and Silurian rocks to the south-west in the
Cargo-Cudal district (Stevens, 1950), to the north in the Wellington district
(Basnett and Colditz, 1946), and closer at hand to the north in the Mumbil
district (Strusz, 1960); intrusions of this type are a likely source for the
dolerite fragments in this formation. The acid volcanic fragments bear a
strong resemblance to the rocks of the Mullions Range Voleanics, part of
which could have been exposed at the time. The fragments in question bear
very little resemblance to the Silurian Canowindra Porphyry (Stevens, 1950;
Ryall, 1965) which outcrops to the west but could be compared with the
early Devonian lavas of the same region, i.e., the Bull’s Camp Volcanics
(Packham and Stevens, 1955), the Duladerry Rhyolite (Stevens, 1954) and
similar rocks in the Cumnock area (Joplin and others, 1952). The last
mentioned is only 12 miles west of Nubrigyn Creek. The quartzite and granite
pebbles are more difficult to account for, they apparently came from further
afield.
The arenites and conglomerates in the vicinity of Nubrigyn Creek are
well sorted (see Table 7) and occasionally cross-bedded. The arenites have
a carbonate cement (Plate x1, fig. 6) or a small proportion of chlorite matrix
(Plate x1, fig. 8). Individual beds are lenticular. These features, as well as
their intimate association with the algal limestones, point to deposition in
a shallow water environment.
To the east of the calcareous facies of the “Nubrigyn Formation” both
arenites and conglomerates are rare and almost all of the succession consists
of slates and silts. Patches of conglomerate have been found in three localities,
viz., two and a half miles west of Euchareena, by the side of the Molong road ;
in the first railway cutting to the north of Euchareena Railway Station; on
the road one and a half miles west of Store Creek Railway Platform. The
152 PALAEOZOIC STRATIGRAPHY AND SEDIMENTARY TECTONICS
first and second localities are important since the conglomerates contain
fossils: Acanthophyllum sp. and Calceola sp. at the first, and Acanthophyllum
sp. and Syringopora flaccida at the second locality. In view of the association
of these conglomerates with fine-grained sediments and their occurrence in
small masses it seems likely that they are slide deposits.
‘South of Nubrigyn Creek, sandy beds are not common in the lower part
of the Cunningham Formation. In the vicinity of Kerr’s Creek, the coarsest
beds are laminated siltstone to very fine sandstone bands. To the east of
this latter area, where the Cunningham Formation is in contact with the
Bay Formation, some of the basal beds of the Cunningham Formation in the
vicinity of the Mullions Range are conglomeratic, containing pebbles of acid
lavas and quartzite. These conglomerates differ from those occurring to the
north-west in the vicinity of the Nubrigyn Limestone in that they contain a
larger proportion of matrix. The arenites and the conglomerates of the
southern area are of the greywacke suite (Plate x1, fig. 5, Table 7) in contrast
to those of the labile sandstone suite in the north-west.
TABLE 7
Micrometric analyses of arenites from the Cunningham Formation in the Huchareena district
A B Cc D
Quartz ash ue Ay aie 21-2 13-8 5-0 3:6
Felspar .. aS a ee 5:1 5:4 10-0 8-8
Igneous rock fragments oe 13-6 36-1 61-5 73-4
Sedimentary rock fragments .. 9-5 5-0 14-7 5-4
Matrixaiine:: ae te a 50:6. 39h 8-1 8-8
A. PT 88. Greywacke, Curragurra Creek, 3 miles east of Euchareena.
B. O120. Greywacke at base of Cunningham Formation, Curragurra Creek. 4 miles
south-east of. Euchareena.
C. O 21. Labile sandstone, from ‘‘ Nubrigyn Formation’’, 4 miles west of
Euchareena.
D. O 29. Labile sardstone, from “‘ Nubrigyn Formation ”’, 73 miles north-west of
Euchareena.
East of Store Creek and Euchareena, arenites are extremely rare. In the
entire section exposed in Curragurra Creek there are only two or three beds
of greywacke (the micrometric analysis of one of these is given in Table 7).
Graded-bedding is common only east of the railway line. The silts
have occasional bands of small-scale cross-bedding and convolute bedding.
Unfortunately most of these structures have been observed in loose blocks
on the sides of hills and it has not been possible to use them as indicators
of the direction of slope of the sea-floor.
A magnificent slump structure is exposed in the narrow gorge of
Curragurra Creek, four miles east of Euchareena. The structure is not so
spectacular as the Ulmarrah Slump in the Hill End district because here
the rock types involved are limited to interbedded silts and slates and the
mass is only about 200 feet thick. The original bedding is still preserved,
normally in folds with axial planes close to the regional dip. In places the
sandy beds are reduced to contorted fragments in a shaly matrix; some of
these fragments of beds are ten or so feet long (Plate x, fig. 8). Examination
of the folds in this slump indicate that the anticlines, recognizable by the
presence of graded bedding in the silts, have their crests facing east, sug-
gesting that the slump has moved towards the east and hence the sea-floor
must have sloped downwards in that direction. This accords with the facies
change of the formation towards shallow-water to the west and establishes
the existence of a trough (the Hill End Trough) of sedimentation to the
east. The eastern margin of this trough has been shown to lie to the east of
= G. H. PACKHAM Lae
the Wiagdon Thrust where shallow water sediments are again developed
the Limekilns area.
It is not known whether this slump is a local phenomenon or whether
it forms an extensive slump sheet. The suggestion was made in the descrip-
tion of slump structures in the Hill End area (p. 66) that they originated
to the west of Hill End and moved eastwards to their present locations.
They could have originated anywhere on the easterly-sloping sea-floor and
thus it may not be possible to trace them back to the western limb of the
Ophir Syncline. The degree to which bedding has been destroyed in the
slumps just to the west of Hill End indicates that they have moved consider-
ably further than the slump in Curragurra Creek.
AGES OF FORMATIONS IN THE EUCHAREENA DISTRICT
The Oakdale and Mumbil Formations of the Euchareena district represent
the southern extension of the two oldest formations of the Mumbil district
(see Table 8). Their occurrence at Mumbil and their fauna have been
described by Strusz (1960, 1961). Although Upper Ordovician graptolites
TABLE 8
Regional correlation table
Quarry Ck.- Mullions Hill End 3 :
Mumbil Eadvale Limekilns
Bis BBGEBESI
Givetian Winburn Tuff
Limekilns Ck. Sh.
Cunningham Formation
Jesse Limestone
Emsian
| Nubrigyn “Fm.” | | Nubrigyn “Fm.” | Fm.”
[eter irre een (et Eeee Teh Tolga Calcarenite Rosedale sua
Siegenian
Sandstn. & congl. Merrions Tuff
ue Bull’s Camp Voles.
Gedinnian Crudine Group
Wallace Cuga Burga Voleanics Bay | IL
; Shale Barnby Cookman Formation
Ludlovian L Hills Mumbil :
Panuara “Fm.” | Shale Fee ROE a ton Chesleigh Formation
masa fee ct a Tia ul ahh Ls.
Wenlockian Bell’s Creek
U tuitions Ra. Voleanics Voleanics
iF Tanwarra Shale
| Panuara “Fm.” | | Panuara “Fm.” | Fm.”
Llandoverian Hy
Malachi’s
Upper Ordovician Hill Oakdale Formation
Sofala Voleanics
Formation
occur in the Oakdale Formation at Mumbil, the highest beds, which contain
limestone lenses, are Upper Ordovician or Lower Silurian on the evidence
of their contained coral fauna. Strusz (1960) considered the overlying
Mumbil Formation to range from “the topmost Llandoverian through most
or all of the Wenlockian’”. Beds which Strusz regarded as basal beds of the
Narragal Limestone (the lower member of the Mumbil Formation) contain
some distinctive forms, including: Palaeophyllum sp., Multisolenia tortuosa
and Acanthohalysites australis. Forms occuring in the main part of the
limestone include: Phaulactis shearsbyi and Entelophyllum latum. Mono-
graptus bohemicus (Lower Ludlow) occurs low in the overlying Barnby
Hills Shale, which is the upper member of the Mumbil Formation. Dr. Strusz
154 PALAEOZOIC STRATIGRAPHY AND SEDIMENTARY TECTONICS
and I recently revisited the area and found that the limestones which he
regarded as basal beds of the Narragal Limestone are separated from it by
a thin succession of acid to intermediate volcanics, and thus the “basal”
limestones are either part of or rest directly on the Oakdale Formation. The
fauna is the same as that in the Oakdale Formation, and has little in common
with the Narragal Limestone. There is no clear evidence then, that the
Narragal Limestone extends far below the top of the Wenlock. In the vicinity
of Euchareena, the Mullions Range Volcanics occur between the Oakdale and
Mumbil Formations. These volcanics can be correlated with the acid to inter-
mediate volcanics which underlie the Mumbil Formation at Mumbil (Table 8).
Further afield at Quarry Creek, 24 miles south-south-west of Euchareena,
the Silurian sequence can be dated with some accuracy by means of the
contained graptolite faunas (Packham, 1968). Beds at Quarry Creek,
containing Monograptus bohemicus and therefore equivalent to the lower
part of the Barnby Hills Shale are underlain by shales containing Upper
Wenlock graptolites (principally M. testis). These are in turn underlain by
cross-bedded sandstones, derived from acid volcanics. Below these sandstones
are Shales and fine quartzitic sandstones containing Upper Llandovery
eraptolites (J. marri is the most common). There is therefore a break in
sedimentation between the Upper Llandovery and the Upper Wenlock, the
result of an uplift called the Quarry Creek Phase of the Benambran Orogeny
by Packham (1967a). Limestone (the Quarry Creek Limestone) underlies
the Upper Llandovery sandstones and shales and overlies Upper Ordovician
andesitic volcanics which can be correlated with the Oakdale Formation.
The clastic succession with limestone bands at the top of what has been
mapped as the Oakdale Formation near Nubrigyn Creek might contain
Llandovery beds and be separate from the volcanics.
In view of the break in sedimentation recognized at Quarry Creek below
the late Wenlock succession it seems likely that the Mumbil Formation ranges
from Upper Wenlock to Ludlow and the Mullions Range Volcanics fall within
the gap in the Quarry Creek succession. The absence of the voleanics at
Quarry Creek and at other localities closer to the Mullions Range implies
that interruption of deposition took place after the extrusion of the volcanics.
The Cuga Burga Voleanics which overlie the Barnby Hills Shale are on
present evidence equivalent to the Bay Formation and have been placed just
above the Silurian-Devonian boundary by Packham (1968) since there is
a considerable thickness of shale above the occurrence of Monograptus
bohemicus in the Barnby Hills Shale. This is a little higher than Strusz
(1960) placed the Cuga Burga Volcanics. The Tolga Calcarenite which over-
lies the Cuga Burga at Mumbil has not been recognized in the area studied,
but it has been found by Wolf (1965) and Kemezys (1959) a few miles to
the north. The latter has found it to be unconformable on the volcanics. The
calcarenite is apparently equivalent to or older than the shale unit in the
Nubrigyn Creek area overlying the volcanics and below the limestones and
lithic arenites. I suggested (Packham, 1967a) that the Tolga Calcarenite
may be equivalent to part of the Garra Formation. This latter unit described
by Strusz (1965) is a succession of shales and limestones over 4,000 feet thick,
forming an outcrop extending about sixty miles north-south and about five
miles east-west. Outcrop commences approximately nine miles west of the
base of the Nubrigyn Formation. I also suggested (Packham, 1968) that
the calcareous “Nubrigyn Formation” because of its considerable content of
terrigenous sands and conglomerates may be younger than the Garra Forma-
tion in which such detritus is rare.
se G. H. PACKHAM 155
Based on his work on the corals, Strusz (1968) has suggested an Emsian
age for the Garra Formation and from the few corals known from the
“Nubrigyn Formation” a correlation with the higher beds of the Garra
Formation would be indicated. Although the conodont faunas of the two
formations are only very sketchily known, no great disparity in age is
apparent. Rare platform conodonts (possibly Polygnathus linguiformis
dehiscens) have been recovered from the highest parts of the Garra Formation
(Philip, 1967). The upper part of the “Nubrigyn Formation” has to date
yielded only small faunas but no platform types have been found. The
conodonts from the lower part of the “Nubrigyn Formation” resemble those
recorded from the lower beds of the Garra Formation by Philip (1967) but
the corals suggest the highest levels. Polygnathus linguiformis foveolata
incidentally, occurs in the Jesse Limestone at Limekilns. It is clear that some
compromise has to be reached. Accordingly, in Table 8, it is suggested (qa)
that the Tolga Calcarenite is equivalent to the base of the Garra Formation,
(6) that the lower part of the Garra Formation may be late Siegenian and
(c) that the upper part is Emsian and equivalent in age to the lower part
at least, of the Nubrigyn Formation but older than the Jesse Limestone.
STRATIGRAPHIC CORRELATIONS THROUGHOUT THE REGION
Although the fossil evidence at present available does not permit precise
dating of the formations, it does, combined with the lithologically established
successions, enable the relative ages of the various stratigraphic units in the
region to be established with some confidence.
Commencing at the base of the succession, the oldest formation on the
east, the Sofala Volcanics may be correlated generally with the Oakdale
Formation, the oldest formation in the west. Whilst Ordovician graptolites
occur in both formations it is possible that the formations might extend into
the early Silurian. In the west there is no definite evidence of Llandovery
sediments but the Tanwarra Shale of the Sofala area is best assigned to the
Upper Llandovery or Lower Wenlock, at the latest. The overlying formation,
the Bell’s Creek Volcanics may then be correlated approximately with the
Mullions Range Voleanics; both are probably Middle Silurian.
The approximate correlation of the Wenlock to Ludlow Mumbil Forma-
tion with the Chesleigh Formation follows from the lithological mapping of
higher formations between Mudgee and Wellington along the Cudgegong
River by Dickson (1962), Shatwell (1962) and Jones (1962). This mapping
has enabled the relationship of the Cuga Burga Volcanics to the Crudine
Group to be established. On the western limb of a iarge anticlinal structure
south of Wuuluman and ten miles east of Wellington, the Cuga Burga
Voleanics are overlain by slates and siltstones and above them is the Merrions
Tuff. In the next anticline to the east, the Cuga Burga Volcanics are not
present but are replaced by tuffs and greywackes of the Turondale Formation.
This relationship indicates that the Turondale Formation and the Cuga Burga
Volcanics can be correlated approximately and so too can the formations
underlying them, the Chesleigh and Mumbil Formations respectively.
No formations in the Euchareena district are at present correlated with
the Waterbeach Formation and the Merrions Tuff. There was apparently
erosion in the west at that time. ;
The Cunningham Formation which is represented in the central and
western parts of the region, passes laterally both eastwards and westwards
into calcareous sediments. The Tolga Calcarenite and at least the lower part
of the “Nubrigyn Formation” are older than the Jesse Limestone judging
156 PALAEOZOIC STRATIGRAPHY AND SEDIMENTARY TECTONICS
by the conodont assemblages (see p. 52). The possibility that the Tolga
Calcarenite and the “Nubrigyn Formation” are equivalent to the Waterbeach
Formation and the Merrions Tuff has not been entirely ruled out. There is,.
however, no clear evidence of calcareous detritus being shed into the western
part of the geosynclinal region even in small quantities from the west until
the. deposition of some 5,000 feet of the Cunningham Formation had taker
place. The highest formation in the east, the Winburn Tuff has no known
equivalent in the central and western parts of the region.
IntTRUSIVE Rocks
This whole region is remarkably free from intrusive bodies compared with
other parts of the Lower Palaeozoic Belt of eastern Australia. Apart from
minor intrusions associated with the andesitic volcanics, the intrusives fall
into two types, viz. granitic masses and dolerite dykes and sills.
Three small, massive, granitic bodies outcrop in the area: the Wiagdon
Granite (named after the Parish of Wiagdon), the Millah Murrah Granite
(named after the Parish of Millah Murrah) and the Bruinbun Granite
(named after a locality on the Macquarie River south of Hill End). In each
case the contacts are sharp and cross-cutting. These granites are probably
part of, or are related to, the Bathurst Granite which outcrops very exten-
sively to the south.
The dolerites occur in two groups roughly symmetrically placed on both
sides of the region. They occur towards the margins of the central strongly
cleaved zone. In the east, they are most common in the upper part of the
Chesleigh Formation. They have been involved in the general low grade
-metamorphism of the region.
STRUCTURE
Folding
Throughout the greater part of the region, the folds have steeply-dipping
axial planes. In the central part of the area a strong cleavage has developed.
This cleavage first appears in the slates and as the degree of deformation
increases it penetrates the coarse rocks as well. Overturned beds occur in
the central part of the area but their dips are steep. It is only in the vicinity
of the Wiagdon Thrust to the south of Wattle Flat that overturned beds
dipping at less than 45° are encountered.
There is a general concordance between the stratigraphy and the struc-
ture of the region. The central area of maximum deposition during Crudine
and Cunningham “times” is the most deformed and has been thrust eastward
over a more stable area. The axial planes of the folds in the western part
of the region dip to the east at angles of 70 to 80 degrees. To the east
dips gradually steepen until at Hill End the axial planes are approximately
vertical; still further to the east they dip steeply to the west. Along the
Turon River, east of Hill End and extending to the north, is a zone in which
the Cunningham Formation has been almost isoclinally folded and the over-
turned limbs of the folds dip steeply to the west. This gives an asymmetry
to the regional structure because there is no corresponding zone to the west
of Hill End. The asymmetry is further emphasized by the thrust faults which
occur to the east. The Hill End anticline marks a very considerable culmina-
tion in the folds of the trough, the sediments exposed along its crest being
some 20,000 feet stratigraphically lower than those in the trough of the Ophir
Syncline to the west and about 15,000 feet below the highest beds in the
isoclinally folded zone of the Cunningham Formation rocks to the east, about
Sally’s Flat.
ZY G. H. PACKHAM 157
The regional cleavage mentioned above seems to be an axial plane
cleavage. It first appears at about the meridian of the Orange-Wellington
railway line and extends eastiwards as far as the Wiagdon Thrust. Although
the rocks of the eastern side of the Wiagdon Thrust, in the area north-east
of Peel, are most strongly overturned, having their axial planes eyping at
20° or so, cleavage is not developed in them.
Apart from the change in dip of the axial planes of the folds there is
also a Swing in strike across the region. In the north the change is from
N 30°W on the western side to N 20°E on the east. The fold axes are closer
packed to the south and they are generally parallel, striking approximately
N 20°W. The plunge is gently towards the north in most folds.
Where grain elongations have been observed and these are common in
the greywackes of the Cunningham Formation, they are approximately
perpendicular to the fold axes and the bedding-cleavage intersections. Pyrite
“shadows” have a similar orientation.
The metamorphism which affected the region was of low grade being
no higher than green-schist facies, though there is an increase in its intensity
from the margins of the area where chlorite, albite and muscovite occur,
towards the centre, where biotite is common. This parallels the stronger
folding and the more intense development of cleavage in the central part of
the region. In addition, accompanying the northerly structural plunge there
is some narrowing of the with of the biotite-bearing zone northwards. Along
the southern margin of the area mapped the zone has a width of about 20
miles while at the latitude of Hill End it is about 17 miles wide.
Faulting
Minor Faults
As far as can be determined at the scale of mapping, faulting is remark-
ably rare in this region. Three minor faults have been mapped; one is a
cross fault in the west of the region near Kerr’s Creek, the second is a high-
angle thrust fault six miles east of Hill End. The third is, in all probability,
also a high angle thrust fault cutting off the Mullions Range Volcanics
along the western side of the Mullions Range. Two localities have been
found where the cleavage has been disturbed by later movements. In neither
case did it seem that faulting significant on the scale of mapping had taken
place. One of these localities is on the Winburndale Rivulet about half a
mile from the Macquarie River and the other on the Macquarie River, one
and a quarter miles upstream from its junction with Lewis Ponds Creek.
The Wiagdon Thrust System
The Wiagdon Thrust fault is the most important tectonic feature of the
region. At present it is known to extend from Gulgong in the north to the
Bathurst Granite near Yetholme in the south, a distance of some eighty
miles. The thrust dips to the west but at varying angles. North of Sofala
the dip is of the order of ten degrees—in some places even less. Near Wiagdon
Hill the dip is considerably steeper and is of the order of sixty degrees. The
strike likewise is very variable. North of Sofala, it is parallel to the fold
axes of the area (N 30°E), and after crossing the Turon River runs approxi-
mately north-south. At Wiagdon Hill, south of Wattle Flat, the fault swings
suddenly towards the south-east and then returns to N10°E and continues
in that direction to Cheshire Creek. This variation in strike, which affects
the folds to the west as well as the fault, appears to be a reflection of the
configuration of the mass against which the sediments to the west were
folded and thrust. In the discussion of the stratigraphy of the Sofala
158 PALABEOZOIC STRATIGRAPHY AND SEDIMENTARY TECTONICS
Voleanics some evidence was presented for the existence of a local volcanic
focus near Sofala in the last stages of the vulcanism.
North of Wiagdon Hill, the thrust is apparently parallel to the bedding
of the over-riding sediments, with the thrust plane occurring at about the
Tanwarra Shale (sometimes within it and sometimes just below it in the
Sofala Volcanics).
To the south at Wiagdon Hill, the thrust occurs within the Sofala
Volecanics and several hundred feet of the volcanics have been thrust over
the Chesleigh Formation. Towards the southern edge of the area mapped the
fault plane passes into the Chesleigh Formation as the result of the develop-
ment of an overturned anticline at the base of the overthrust mass. Tiwo
minor thrusts occur on the foreland side in the south.
Instead of the Sofala Volcanics being swept clean of later formations
on the foreland side by the thrust, south-east of Wiagdon Hill the entire
sequence has been preserved. This is the direct result of a southerly plunge
of the folds on the foreland side which may be attributed indirectly to the
suggested thinning of the volcanic pile at the top of the Sofala Volcanics.
Just to the east of the thrust in this southern area the greatest degree of
overturning in the whole area has been observed. Overturned dips between
65° and 20° are normal. In exceptional cases the beds are horizontal (over-
turned) and in one local pucker a dip of 40° to the east was observed; this
overturned zone is bounded by a minor thrust which dies out to the north
where the main thrust swings to the west.
North of the Turon River, the thrust structures again become complex.
About a mile north of the river a new dislocation becomes apparent, this
time to the west of the main thrust line. It appears within the Chesleigh
Formation bringing the upper part of the Chesleigh Formation against the
Bell’s Creek Voleanics and then, further north, the Cookman Formation
against the Bell’s Creek Voleanics. The main Wiagdon Thrust is again in
the Tanwarra Shale or just within the Sofala Volcanics.
The Wiagdon Thrust can be observed where it crosses the Turon River
and at a number of localities about five miles north of Sofala in tributaries
of Cookman Creek. In these planes there is a transitional zone from sheared
andesite of the Sofala Volcanics to phyllite derived from the Tanwarra Shale.
The original bedding of the Tanwarra Shale is broken down but deformed
traces of it remain both on the microscopic and the macroscopic scale.
Razorback Thrust
This fault was discovered by Day (1961) four miles north of the Turon
River in the vicinity of the Razorback Mine from which it takes its name.
This fault is on the extreme east of the area and brings the Sofala Voleanics
up against westerly-dipping Upper Devonian sandstones cutting off the
western limb of the synclinal structure in which the sandstones lie. There
is sufficient relief in the vicinity of the Turon River to determine that the
fault plane dips steeply to the west. It is the only structure in the area for
which a post-Upper Devonian age can be demonstrated.
GENERAL SUMMARY OF DEPOSITIONAL HISTORY
Ordovician
Ordovician rocks are exposed on the eastern and western margins of the
region and andesitic volcanic material dominates both areas. The few sedi-
mentary structures observed in the lower part of the Sofala Voleanics suggest
an eastward slope of the sea-floor and the deposits represent a turbidite facies.
e G. H. PACKHAM 159
Higher, pyroclastic debris becomes more abundant and the presence of occa-
sional limestone lenses in the upper part of the formation implies some
shallowing but this may be only of local significance. The stratigraphy of
the Oakdale Formation is unknown in detail but limestone lenses occur within
the higher parts of the Upper Ordovician in the formation and at lower strati-
graphic levels further west (e.g., at Molong and Bowan Park). This evidence,
though fragmentary is consistent with the palaeogeographic pattern which
existed in the later Silurian. The sea-floor sloped generally to the east into
a region of turbidite deposition (the Hill End Trough) and commencing at
the western margin of the region under discussion and extending further
west, was an elevated zone (the Molong Geanticline). A second trough of
turbidite deposition, the Cowra Trough, was probably in existence at this
time west of the geanticline (Packham, 1967).
WwW Kerr’s =
Creek ; Hill End Sofala
LEGEND
Def Cunnin gham Formation
Dmt Merrions, Tuff
Dwf Waterbeach Formation
Dtf Turondale Formation
Sem Cookman Formation
Scf Chesleigh Formation >]Osv Sofala Voleanics
Smf Mumbil Formation ig Oo Oakdale Formation © 2 a enimis.
Sbv Bell's Creek Volcanics @
Stw Tanwarra Shole ©
Smv Mullions Range Volcanics
SILURIAN DEVONIAN
Figure 10. Cross-section from Sofala to Kerr’s Creek. Section projected to depth
to demonstrate stratigraphic relationship between formations in eastern and western
margins of the Hill End Trough.
Silurian
It is not known whether the Sofala Voleanics and the Oakdale Formation
extend into the Silurian. The possibility exists that deposition may have
been interrupted by early phases of the Benambran Orogeny. The Tanwarra
Shale though not definitely proven to be Lower Silurian, is probably of that
age or basal Wenlock at the latest. The conglomerate at the base of the
formation represents an erosional break which might be correlated with one
of the two Lower Silurian orogenic phases of the Benambran Orogeny in the
Orange district, the Cobbler’s Creek Phase at about the base of the Llandovery
or the Panuara Phase at about the middle of the Llandovery (Packham,
1968). If the correlation can be made the later one is the more likely.
The two acid volcanic formations which follow, the Mullions Range
Volcanics in the west and the Bell’s Creek Volcanics in the east are apparently
Middle Silurian. It was suggested that the Mullions Range Volecanics lay
within the gap in deposition from late Llandovery to Upper Wenlock found
to occur at Quarry Creek. Mapping of the base of the volcanics has failed
to reveal the presence of Llandovery beds beneath them except perhaps in
the north, giving the impression that there is at least a disconformity
between the Mullions Range Volcanics and the Oakdale Formation. Because
of the absence of the Mullions Range Volcanics in nearby sections to the
west, such as the one along the Bell River only four miles west of the 1,500
feet-thick type section of the volcanics, it is likely that the voleanics antedate
the Quarry Creek Phase of the Benambran Orogeny. The regional distribu-
tion of the Bell’s Creek Volcanics is also suggestive of tectonic movements
post-dating them. The Chesleigh Formation which normally overlies the
Bell’s Creek Volcanics, overlies the Sofala Volcanics in the vicinity of Wattle
160 PALAEOZOIC STRATIGRAPHY AND SEDIMENTARY TECTONICS
Flat, east of the Wiagdon Thrust. If these relations are the result of a
local uplift, the uplift is roughly contemporaneous with the Quarry Creek
Orogenic Phase.
The lower parts of the Chesleigh Formation and the Barnby Hills Shale
are composed of sediments consisting of quartz, white mica and some chlorite;
volcanic debris is rare. Deposits of this composition are common in Ludlow
successions in the southern highlands of New South Wales, and sedimentary
structures in the Chesleigh Formation in the Sofala area indicate a down-
ward slope of the sea-floor to the east-north-east. The detritus was probably
derived from Ordovician sediments (with minor contributions from granitic
sources), exposed in the southern highlands of New South Wales and eastern
Victoria as a result of the Benambran Orogeny. The sediments might have
been transported northwards along the trend of the Benambran structures and
thence into the trough. The sediments deposited on the Molong Geanticline
at this stage, apart from the Narragal Limestone and its equivalents, are
fine-grained (siltstones and shales) although their environment of deposition
is unknown. The sediments in the Hill End Trough are coarser, consisting
of interbedded slates, siltstones and fine-grained greywackes and having the
typical features of a turbidite facies.
The dacitic volcanic debris in the upper part of the Chesleigh Formation,
which is intermingled with detritus similar to that in the lower part, has
no known equivalent in the Euchareena area. It occurs in both the Hill End
and Sofala occurrences of the formation, though in the latter it is a minor
constituent south of the Turon River, increasing rapidly in abundance north-
wards. Deposition throughout is in turbidite facies and the directional
features of the sedimentary structures are consistent with those of the lower
part of the formation. The volcanic centre supplying the detritus probably
lay to the west or south-west of Hill End. It has not as yet been identified.
The highest formation regarded as Silurian in the Sofala area is the
Cookman Formation. At its base, an important change in the pattern of
sedimentation took place. Sedimentary structures indicate a reversal of the
direction of source of the sediments. From this formation onwards, in the
Hill End and Sofala areas, detritus came from the east. The Cookman
Formation is a sequence of slates and quartz-rich greywackes. Their petro-
graphy is consistent with their having been derived from the Chesleigh
Formation with minor contributions from underlying formations. The uplift
of the structure to the east (the Capertee Geanticline) which considerably
restricted the Hill End Trough can be regarded as part of the Bowning
Orogeny and may be contemporaneous with the Yarralumla Phase (Opik
1958). The upper part of the Barnby Hills Shale was probably deposited
at the same time as the Cookman Formation.
Devonian
The new palaeogeographic pattern established with the commencement
of deposition of the Cookman Formation persisted throughout the remainder
of the history of the Hill End Trough. At approximately the beginning of
the Devonian, vulcanism was renewed, this time on a grand scale, especially
in the east. After the basal breccias of the Crudine Group, the remainder
of the Turondale Formation consists of thickly bedded dacitic tuffs, grey-
wackes composed of tuffaceous material, conglomerates and interbedded slates
and siltstones. The Cuga Burga Volcanics were poured out in the northern
part of the Euchareena area and northwards extending down into the Hill
End Trough. Dacitic vuleanism responsible for the Bay Formation produced
clastic sediments similar to those of the Turondale Formation in the southern
f G. H. PACKHAM 161
part of the Euchareena area at about this time. The axis of the trough lay
west of the Hill End Anticline as it must have done during the deposition
of the Cookman Formation.
The Waterbeach Formation (the upper part of the Crudine Group) is
free from evidence of contemporaneous volcanic activity. Nevertheless its
greywackes are composed almost entirely of material ultimately of volcanic
origin and similar to that of the Turondale Formation. Except for the sedi-
ment contained in the Ulmarrah Slump which is the most westerly occurrence
of the formation, the detritus appears to have been derived from the east.
The axis of the trough remained just west of the Hill End Anticline.
Wolf (1965) and Kemezys (1959) have mapped an erosional junction at
_ the top of the Cuga Burga Volcanics and the distribution of the Cuga Burga
Volcanics and the Bay Formation in the Euchareena area is compatible with
such a contact. Any uplift on the western side of the trough would have
probably increased the slope on the eastern side of the Molong Geanticline
and initiated the Ulmarrah Slump.
The axis of the trough seems to have been at its western limit during
the deposition of the Merrions Tuff, lying at least eight miles west of the
Hill End Anticline and less than 16 miles east of the present outcrop of the
Oakdale Formation. The thick beds of dacitic pyroclastic material of the
Merrions Tuff, like those of the Turondale Formation have been interpreted
as some kind of turbidite deposit. While it is possible that turbidites could
have flowed up the western side of the trough to some extent, the distance
was probably not significant, especially since dacite flows within the formation
are found west of the Hill End Anticline. The westward shift of the axis
may have been the result of infilling from the east.
Vulcanism in the trough ended with the Merrions Tuff, and with the
return of normal greywacke sedimentation in the Cunningham Formation,
the axis of the trough moved eastwards to about the Hill End Anticline.
The sediments of the central part of the trough are fine but in the east pass
into a zone in which greywackes and conglomerates are common. In some
of these beds the amount of matrix is small. This distribution suggests that
the slope was short and perhaps steep and that the main part of the trough
was flat. The slope on the west may have been more regular since slump |
structures are common as far east as Hill End. The Limekilns Group which
overlies the Merrions Tuff, east of the Wiagdon Thrust is correlated with the
Cunningham Formation. The age of the limestone in the middle of the Group is
Emsian to perhaps basal Eifelian on the basis of its conodont fauna. Judging
by its fauna the Limekilns Group was deposited largely in the neritic
environment. No indication of turbidite deposition has been found in it. On
the west, the Cunningham Formation passes into the calcareous facies of the
“Nubrigyn Formation”. The transition is extremely rapid. The “Nubrigyn
Formation” apparently represents a near-shore turbulent environment with
algal reefs, limestone breccias, abundant coarse terrigenous material
derived from a variety of sources with evidence of local vulcanism. These
pass eastwards rapidly into sequences of slates and siltstones with rare
greywackes, boulder horizons and calcareous turbidite beds. The basal part,
if not all, of the “Nubrigyn Formation” is older than the Jesse Limestone.
Overlying the Limekilns Group in the vicinity of Limekilns and extending
south for some miles is the Winburn Tuff, a formation which has lithological
features very similar to those of the Merrions Tuff but is much richer in
orthoclase. The field relations are strongly suggestive of a unconformity
with the Limekilns Group. The Winburn Tuff is in turn overlain unconform-
ably by quartzites and shales of the Upper Devonian Lambie Group. The
KE
162 PALAEOZOIC STRATIGRAPHY AND SEDIMENTARY TECTONICS
latter unconformity is representative of a phase of the Tabberabberan Orogeny.
The former might be regarded as related to an earlier phase of the same
orogeny.
Acknowledgements
I am grateful to Associate Professor T. G. Vallance for critically reading
the-manuscript of this paper and making many helpful suggestions. I would
also like to thank Miss J. Forsyth and Miss P. L. Vockenson for drafting
the figures and Mr. G. Z. Foldvary for printing the photographs and
assembling the plates. Field expenses incurred during this work were met
from a Sydney University Research Grant and funds from the Australian
Research Grants Committee.
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Harprr, L. F., 1918.—The Hill End and Tamboroora goldfield. N.S.W. geol. Surv. min.
IAGIB > CA
JONES, J. G., 1962.—The geology of the parishes of Wuuluman and Yarragal. Unpubl.
B.Sc. hons. thesis (Sydney University).
Jounson, J. H., 1964——Lower Devonian algae and encrusting foraminifera from New
South Wales. J. Palaeont., 38: 98.
JOPLIN, and OTHERS, 1952.—A note on the stratigraphy and structure of the Wellington-
Molong-Canowindra region. Proc. Linn. Soc. N.S.W., 77: 83.
Joptinc, A. V., 1950.—The geology and ore deposits of the Hill End-Tamboroora district,
N.S.W. Unpubl. M.Se. Qualifying thesis (Sydney University).
Kemezys, K. J., 1959—Geology around Bakers Swamp. Unpubl. B.Sc. hons. thesis
(Sydney University).
Opix, A. A., 1958—The geology of the Canberra district. Bull. Bur. Miner. Resour. Geol.
Geophys. Aust.: 32.
PackHAm, G. H., 1958.—Stratigraphic studies in the older Palaeozoic rocks of the
Tasman Geosyncline in central-western New South Wales. Unpubl. Ph.D. thesis
(Sydney University).
, 1960.—Sedimentary history of part of the Tasman Geosyncline in south-eastern
Australia. Int. Geol. Congr., 21(12): 74. ;
, 1962.—An outline of the geology of New South Wales. 36th Meeting Austr.
& N.Z. Assoc. Adv. Sci., Sydney. A Goodly Heritage: 24.
, 1967.—The occurrence of shelly Ordovician strata near Forbes, N.S.W. Aust.
J. Sci:z, 30: 106-107.
, 1968.—(Ed.) Geology of New South Wales. J. geol. Soc. Aust.: 17 (in press).
Puitip, G. M., 1967.—Stratigraphic correlation of some of the principal Devonian lime-
stone sequences of eastern Australia. Proc. int. Symp. Devonian System, Alberta
Soc. Petroleum Geologists, Calgary (in press).
Pittman, E. F., 1881.—Geological map of Hill End of Tamboroora. Ann. Rept. Dept.
Mines N.S.W. (1879).
Porter, P. E., and Siever, R., 1955.—A comparative study of Upper Chester and Lower
Pennsylvanian stratigraphy variability. J. Geol., 63: 439.
Raw, F., 1943.—Some altered palagonite tuffs from Jamaica and the origin and history
of their chlorites. J. Geol., 51: 215.
Ryatu, W. R., 1965.—The geology of the Canowindra East area, N.S.W. J. Proc. r. Soe.
N.S.W., 98: 169-179.
SAnpDerS, J. E., 1965——Primary sedimentary structures formed by turbidity currents
and related resedimentation mechanisms. Spec. Publ. Soc. Econ. Palaeont. & Mineral.
12: 192-219.
SHATWELL, D. O., 1962.—The geology of the parishes of Merinda, Rouse, Wiadere and
Piambong, near Mudgee, N.S.W. Unpubl. B.Sc. hons. thesis (Sydney University).
\
Proc. LINN. Sac
me, y “g
e.
BGG
The Lower and Middle Palaeozoic stratigraphy and Sedimentary Tectonics of the
Sofala-Hill End—Huchareena region, N.S.W.
W uous
Di ee
ei
Proc. Linn. Soc- N.S.W., Vol. 93, Part 1 PLATE X
The Lower and Middle Palaeozoic stratigraphy and Sedimentary Tectonics of the
Sofala-Hill End—Euchareena region, N.S.W.
Proc. Linn. Soc. N.S.W., Vol. 93, Part 1 Piatp XU
GEOLOGICAL MAP OF THE EUCHAREENA-
HILL END- SOFALA AREA
, o ’ 2 z
——
Scale in Miles
Bieta
yy Yy Se
YY
TERTIARY
vpPER
DEVONWAN
SS
=
SH
LOWER TO
MIDDLE
DEVONIAN
SILURIAN
Sala Voloarics
Oakdale Formation
Lumestones in Mumbii Formation Ls
IRDOVICLAN |
Direction of slope of sea Floor
indicated by sedimentary structures
Moderate to low angle thrust faults Jf
6
Other Faults ff
: Wiagdon Grerite
Dip of vedding |} Overturned dip INTRUSIVES
Roads Railway line
G. H. PACKHAM 163
Suerrarp, K. M., 1954.—The assemblages of graptolites in New South Wales. J. Proc. r.
Soc. N.S.W., 87: 73.
Stevens, N. C., 1950.—The geology of the Canowindra district, N.S.W. Pt. 1. The
stratigraphy of the Cargo-Toogony district. J. Proc. r. Soc. N.S.W., 82: 319.
, 1954.—Studies in the Palaeozoic geology of central-western N.S.W. Unpubl.
Ph.D. thesis (Sydney University).
Strusz, D. L., 1960—The geology of the parish of Mumbil, near Wellington, N.S.W.
J. Proc. r. Soc. N.S.W., 93: 127.
, 1961.—Lower Palaeozoic corals from New South Wales. Palaeontology, 4: 331.
, 1965 —A note on the stratigraphy of the Devonian Garra Beds of New South
Wales. J. Proc. r. Soc. N.S.W., 98: 85.
—, 1968.—The development of the Molong Geanticline in the Devonian of New
South Wales. Proc. Int. Symp. Devonian System, Alberta Soc. Petroleum Geologists,
Calgary (in press).
TYRRELL, G. W., and Pracock, M. A., 1926.—The petrology of Iceland. Trans. r. Soc.
Edinburgh, 55: 51. ;
Wotr, K. H., 1965.—Petrogenesis and paleoenvironment of Devonian algal limestones
of New South Wales. Sedimentology, 4: 113.
Wricut, A. J. T., 1966.—Studies in the Devonian of the Mudgee district, New South
Wales. Unpubl. Ph.D. thesis (Sydney University).
EXPLANATION OF PLATES
PLATE Ix
Fig. 1. Breccia composed of fine-grained sedimentary rocks, some containing quartz
veins. x8. Sofala Volcanics, Turon River, Portion 53 Parish of Stewart. Fig. 2.
Subgreywacke. x8. Chesleigh Formation, upper part of Wiagdon Hill, 23 miles south
of Wattle Flat. Fig. 3. Chloritic patch in tuffaceous arenite. x 8. Turondale Formation,
14 miles south-east of Turondale. Fig. 4. Subgreywacke. x8. Cookman Formation, Type
section of formation, Turon River, 34 miles west of Sofala. Fig. 5. Tuffaceous arenite.
x8. Turondale Formation, Type-section, Turon River, 1,300 feet above the base of the
formation. Fig. 6.Greywacke from the base of bed 80” thick. x8. Waterbeach Forma-
tion, Type-section, Turon River, 1,350 feet above the base of the formation. Fig. 7.
Greywacke from same bed as above. 70’ above the base of the bed. x8. Fig. 8. Load
casts on the base of a subgreywacke. Chesleigh Formation, 3% miles west-north-west
of Limekilns.
PLATE X
Fig. 1. Graded-bedded silts and slates. The photograph represents three feet of
section. Waterbeach Formation, junction of the Macquarie River and Winburndale
Rivulet. Fig.-2. Greywacke conglomerate with a large proportion of matrix. Turondale
Formation, Turon River, 2 miles north of Turondale. Fig. 3. Graded-bedding in silts
and slates in overthrust beds east of the Wiagdon Thrust. Crudine Group, 3 miles north-
west of Limekilns. Fig. 4. Coarse breccia containing blocks of dacite. Merrions Tuff,
Type section, Turon River, 1,550 feet above the base of the formation. Fig. 5. Tuffaceous
arenite with chloritic patches. Merrions Tuff, 2 miles north of Limekilns. Fig. 6.
Outcrop of mudflow horizon (the hammer rests on sandy phase) showing contact
between sandy and muddy phases. Cunningham Formation, Type section, Turon River,
2,150 feet above the base of the formation. Fig. 7. Boulder bearing greywacke injected
into slates or siltstones in the Ulmarrah Slump. Waterbeach Formation, Ulmarrah
Anticline, Pyramul Creek. Fig. 8. Contorted siltstone band in slump structures.
Cunningham Formation, Curragurra Creek, 5 miles north-north-east of Huchareena.
PLATE XI
Fig. 1. Tuffaceous arenite containing abundant epidote. x8. Merrions Tuff, Type
section, Turon River, 850 feet above the base of the formation. Fig. 2. Matrix of mudflow
horizon. x8. Cunningham Formation, Type section, Turon River, 2,150 feet above the
base of the formation. Fig. 3. Greywacke. x8. Cunningham Formation, ? mile west of
eastern boundary of the formation on the Sofala-Sally’s Flat road. Fig. 4. Greywacke.
x8. Cunningham Formation, Type section, Turon River, 2,500 feet above the base of the
formation. Fig. 5.Greywacke. x8. Base of Cunningham Formation, Curragurra Creek,
4 miles south-east of Euchareena. Fig. 6. Labile sandstone with carbonate cement. ~x 8.
“Nubrigyn Formation’, 4 miles west of Euchareena. Fig. 7. Greywacke matrix of the
- Ulmarrah Slump. Waterbeach Formation, Ulmarrah Anticline Pyramul Creek. Fig. 8.
Labile sandstone. x8. “Nubrigyn Formation”, 4 miles west of Euchareena.
PLATE XII
Geological Map of the Sofala-Hill End-Euchareena Region.
K K
THE CONSTITUTION, DISTRIBUTION AND RELATIONSHIPS
OF THE AUSTRALIAN DECAPOD CRUSTACEA
A PRELIMINARY REVIEW*
D. J. G. Grirrin and J. C. YALDWYN
The Australian Museum, Sydney, N.S.W.
[Read 24th April, 1968]
Synopsis
Approximately 1200 species are recorded from Australia, representing 57 of the
73 known decapod families. These are placed in 361 genera. About 54% of the
species belong to the Brachyura. Xanthid and majid crabs as well as parastacid
crayfish are well represented. Little is known of many natant groups. The majority
of species are tropical and many are widespread northern forms; there are a smaller
number of widespread southern forms. The tropical fauna shows close affinities with
the faunas of other Indo-West Pacific areas.
INTRODUCTION
There are approximately 1,200 species of Crustacea Decapoda currently
recognized and recorded from the Australian area representing 57 of the 73
families into which the living members of this order are at present divided. The
Australian species range in size from the transparent, planktonic, sergestid
shrimp Lucifer, a few mm. in length, through the whole range of variously-
sized prawns, crayfish, hermit crabs and true crabs, up to the giant xanthid
crab of the southern continental shelf, Pseudocarcinus gigas, reaching at
least 30 lb. in weight and 13 inches across the carapace. Decapods are found
on land, in freshwater, on sandy beaches, in mangrove swamps and in the
sea, from the surface of the open ocean through all depths, and on all
bottoms, down to the floor of the nearby hadal trenches of this area. On the
continent itself, species occur from the upper slopes of the Snowy Mountains,
through the streams and swamps of almost all land and vegetation types, to
the intermittent waters of some parts of the inland deserts.
The present review is an attempt to systematize certain aspects of the
currently available knowledge of the diversity, distribution and relationships
of the Australian Decapod fauna.
The data used in this review come from a series of largely uncritical
family and subfamily checklists prepared from the decapod literature index
held in the Crustacea Department at the Australian Museum. This index
was initiated by A. R. McCulloch early this century, but has been greatly
expanded and developed by F. A. McNeill since then. McNeill in fact, since
his retirement as Curator of Crustacea in 1961, has still remained largely
responsible for keeping this unique and invaluable index up to date. Without
its help we would have been unable to present this study in its present form.
The index is basically a cross-referenced file of decapod species recorded in
published literature for Australia and nearby areas. It pays special regard
to synonymy, recorded localities and illustrations. It also contains numerous
unpublished observations, new records, extensions to range, suspected
* The revised text of a paper presented at the Australian/New Zealand Meeting
on Decapod Crustacea, Sydney, October 24-28, 1967.
PROCEEDINGS OF THE LINNEAN Soormry or New SoutH WALES, Vor. 93, Part 1
D. J. GRIFFIN AND J. C. YALDWYN 165
synonyms, drawings, photographs and colour sketches. Though incomplete,
its imperfections are largely known, and it is being continually expanded
and developed. There are at present approximately 28,000 items individually
indexed and systematically arranged in this file.
The present review is restricted to Australian species and localities
actually published, in press or in draft manuscript known to us in July, 1967.
In a very few cases certain clear, specific omissions or new state records,
known to several workers on our fauna, and not in the above categories, are
included. Many new records for Australia, recorded only in the Museum
literature file, or known only to workers with an Australian group under
review, are not included in the present study. No attempt has been made
to search the collections of the Australian Museum, or other State museums,
for extensions of range or to confirm the presence or absence of species from
areas considered in the tables below. Such a search was not considered neces-
sary or practicable at this stage; in the opinion of the authors, the extra
details obtained would not alter to any significant extent the overall con-
clusions about relationships and distribution patterns drawn here.
The classification used at the suprageneric level in this review is mainly
that set out in Balss (1957), though this has been modified in some instances
to suit our views. For example, Holthuis (1955) is followed for the caridean
shrimps and prawns, Griffin (1966) for the majid spider crabs and Stephenson
and Campbell (1960) for the portunid swimming crabs.
HISTORICAL
The only previous attempt to review the Australian decapod Crustacea
was made by Haswell in 1882. His well-known “Catalogue of the Australian
Stalk- and Sessile-Eyed Crustacea” was published by the Australian Museum
as the first of a series of similar catalogues on various groups of the
Australian fauna. Haswell’s section on the Decapoda in this volume includes
381 species representing 152 genera. These genera are distributed among 37
families or family equivalents. Geographical information is very sparse and
usually restricted to one or two individual localities for each species; no
summary or distribution analysis is included in this work. Assuming that
all the species recorded by Haswell still stand in the Australian list, the 1882
Catalogue covers only 32% of the decapods now known from this area.
Since Haswell’s major work on this order, a number of family and
generic reviews have appeared, covering either the whole of Australia or a
restricted area. A number of important expedition reports, covering usually
one major collection for a restricted area, have also been issued in this
period. Only three groups of Australian decapods are considered to be
adequately covered by modern systematic reviews: the penaeid prawns (Dall,
1957; Racek and Dall, 1965), the majid spider crabs (Griffin, 1966) and the
portunid swimming crabs (Rees and Stephenson, 1966; Stephenson, 1961,
1962; Stephenson and Campbell, 1959, 1960; Stephenson and Hudson, 1957:
Stephenson, Hudson and Campbell, 1957). Other published systematic and
geographic accounts of major significance are as follows: freshwater par-
astacid crayfish (Clark, 1936, 1941; Riek, 1956); parastacid crayfish of
Queensland, of Western Australia and of Tasmania (Riek, 1951b, 1967,
1967b) ; porcellanid crabs of Western Australia (Haig, 1965) ; oxystome and
gymnopleuran crabs of Western Australia (Tyndale-Biscoe and George, 1962) ;
freshwater potamid crabs and the zoogeography of freshwater decapods in
general (Bishop, 1963, 1967); Ocypode crabs of Western Australia (George
and Knott, 1965), and an illustrated account of the decapods of South
166 THE AUSTRALIAN DECAPOD CRUSTACEA
Australia (Hale, 1927). Expedition reports of importance are as follows:
Abrolhos Islands, Western Australia (Percy Sladden Trust Expedition—
Montgomery, 1981) ; Low Isles and adjacent reefs, Queensland (Great Barrier
Reef Expedition—MeNeill, 1968); Port Curtis district, Queensland (Grant
and McCulloch, 1906) ; continental shelf off Sydney, New South Wales (Thetis
Expedition—Whitelegge, 1900); continental shelf off south-eastern and
southern Australia (Hndeavour Expedition—Rathbun, 1918, 1923; Schmitt,
1926); shelf and deep waters off Tasmania and southern Australia
(B.A.N.Z.A.R. Expedition—Hale, 1941), and south-western Australia
(Hamburg Museum Expedition—Balss, 1935).
Siz—E AND CONSTITUTION OF THE FAUNA
The 1200 or so species of decapod crustaceans recorded from Australia
belong to 361 genera and 57 families. The largest systematic group is the
Brachyura or true crabs which includes about 54% of the total species
(Table 1). The smallest group is the Macrura Reptantia (crayfish and allies)
which contains almost as many species as the Anomura or hermit crabs but
only half the number of genera; the Natantia (shrimps and prawns) comprise
less than three times as many species as the reptant macrurans, but spread
through almost four times as many genera.
TABLE 1
Size of the four major systematic groupings in the Australian decapod fauna
Number Number Number Percentage
Group a of of of of Total
Families Genera Species Species
Natantia .. Shrimps and prawns 16 82 299 25-1
Macrura
_ Reptantia Crayfish, ete. 4 2) 119 10-0
Anomura .. Hermit crabs, etc. 14 41 129 10-8
Brachyura True crabs 23 217 643 54-1
Total are ae ae af 57 361 1190 100-0
The largest family is the Xanthidae with 47 genera and 166 species
(Table 2). The largest genera are the xanthids Actaea and Pilumnus with
27 species each. The Parastacidae with only 11 genera contains at least
91 species; included are three particularly large genera, Cherax, EHuastacus
and Hngaeus. The average number of genera per family is 6:3, the average
number of species per family is 21 and the average number of species per
genus is 3:3.
Considering the larger families and genera (Table 2) we find that 17
of the families (30% of the familial total) contain more than 5 genera (in
all 278 genera—75% of the generic total), that 23 families (421% of the
familial total) contain 998 species (83-9% of the specific total) and that
253 genera (64% of the generic total) account for 365 species (30:-4% of the
specific total). In these larger groups then, the average number of genera
per family is 16, the average number of species per family is 45 and the
average number of species per genus is 16.
Among the best represented groups of decapods in Australia are the
Parastacidae (freshwater crayfish)—probably about 80% of the total world
species; the Mictyridae (soldier crabs)—all three species are represented
in Australia, and the anomuran family, the Lomisidae (one species), which
a D. J. GRIFFIN AND J. C. YALDWYN 167
is confined to Australia. The intertidal grapsid crabs, although one of the
larger families (4% of the total marine species, Table 5), is in fact rather
poorly represented in Australia—less than 25% of the total known species
occur here; particularly noticeable is the paucity of varunines. The terrestrial
groups—Gecarcinidae (land crabs) and Coenobitidae (robber crabs and land
hermits )—are very poorly represented with only three species out of ‘a world
total of about 15. Several families of natants are poorly represented in the
Australian fauna. Particularly noticeable is the paucity of pandalid and
crangonid shrimps. These families are especially well represented in northern
hemisphere waters. The Australian fauna includes at present only six
pandalids (world total about 112) and 10 crangonids (world total about
TABLE 2
List of the large families (containing more than 10 genera) and genera (containing 10 or more species)
No. No. No. No.
Family of of Genus of Genus of
Genera Species Species Species
20 100 Xanthidae 47 166 Actaea 27 +Pilumnus 27
or more SPecies: Carpilodes 10 Xantho 10
ees Majidae 43 93 Hyastenus 13
Parastacidae 11 91 Cherax 24 Euastacus 19
Engaeus 15
50-99 Thalamita. 23 Portunus 18
species Portunidae 15 80 Charybdis 14
11-19 Leucosiidae 19 74 Leucosia 15 Kbalia 10
genera ' Paguridae 15 58 Pagurus 12
Penaeidae 15 50 Metapenaeopsis 10
Palaemonidae 19 48 Macrobrachium 12
Grapsidae 17 43 Sesarma 15
Ocypodidae 11 41 Macrophthalmus 15 Ueca 13
Ls 20-49 Goneplacidae 14 21
species Porcellanidae 8 38 Petrolisthes 10
: Alpheidae 6 31 Alpheus 17
6-10 _ Parthenopidae 10 29 Parthenope 16
genera Dromiidae 9 24
Hippolytidae 9 19
pa) Atyidae 6 19
Scyllaridae 5 15 Seyllarus 10
Calappidae 3 14
10-19 Pinnotheridae 3 12
5 species Palinuridae 4 11
genera Galatheidae 3 11
or less Hymenosomidae 3 10
140). A number of other natant families—stenopodids, sergestids and
processids—present in Australia in small numbers, conform to a general
world familial pattern of a few species, often with a limited geographical
distribution, in each major area. Less than 1/5 of the known Atyidae (fresh-
water shrimps) and less than 1/10 of the Potamidae (freshwater crabs) occur
in Australia—this contrasts with the strong representation of the other
freshwater family, the parastacid crayfish.
Masor Hapsrrat GROUPINGS
The vast majority of the species (90%) are exclusively marine. However,
all of the four major systematic groupings are represented on land or in
freshwater. The list below gives the names of the families (and in some
cases genera) occurring in freshwater, on land and in the various zones
of the sea. Three families, including 18 genera and more than 116 species,
168 THE AUSTRALIAN DECAPOD CRUSTACEA
are confined to freshwater and four other genera (of three families) also
occur in freshwater. Two genera containing four species belonging to two
families are found on land.
The intertidal zone is dominated by representatives of five families which
do not normally occur subtidally (a total of more than 100 species). Thirteen
other families are strongly represented intertidally or immediately sub-
tidally ; the most important of these are the xanthids, portunids, majids and
leucosiids.
Five families contain mainly pelagic or bathypelagic oceanic species and
the representatives of 20 families are characteristic of the shelf. Particularly
important shelf groups are the majids, parthenopids, leucosiids, portunids
and dromiids. A list of these habitat groupings is given below.
Freshwater Families and Genera
Restricted freshwater families: Atyidae, Parastacidae, Potamidae.
Freshwater genera or species: Macrobrachium and Palaemonetes (in
part) (Palaemonidae) ; Halicarcinus lacustris (Hymenosomidae) ; Varwna
litterata (partly) (Grapsidae). ;
Terrestrial Families
Restricted terrestrial families: Coenobitidae, Gecarcinidae.
TAttoral (i.e., Intertidal) Families
Restricted littoral families: Thalassinidae,.-Lomisidae, Mictyridae,.
Grapsidae, Ocypodidae.
Other families with some littoral (but not exclusively littoral) represen-
tatives: Palaemonidae, Alpheidae, Gnathophyllidae; Hippolytidae, Steno-
podidae, Axiidae, Callianassidae, Dromiidae, Calappidae, Leucosiidae, Majidae,
Portunidae, Xanthidae.
Oceanic Families
Restricted pelagic or bathypelagic families: Penaeidae (in part only),
Sergestidae, Oplophoridae, Pasiphaeidae, Grapsidae (one genus only—Planes).
Families characteristic of Shelf and Shallow Waters
Penaeidae, Ogyrididae, Stenopodidae, Palinuridae, Scyllaridae,
Laomediidae, Galatheidae, Albuneidae, Dromiidae, Raninidae, Calappidae,
Homolidae, Leucosiidae, Majidae, Parthenopidae, Hymenosomidae, Portunidae,
Xanthidae, Goneplacidae, Cancridae.
COMMERCIAL GROUPS
The commercial species are largely macrurous—natant prawns and
reptant crayfish. The major families are the Penaeidae, Palinuridae and
Portunidae (part only) with 13 genera and about 50 species being fished
commercially. The Scylaridae and Parastacidae are of minor economic
importance.
GEOGRAPHICAL DISTRIBUTION WITHIN AUSTRALIA
The main feature of the patterns of geographical distribution of the
marine decapods in Australia is the fairly clear partitioning of the fauna
into a northern (tropical) group and a southern (temperate) group, the
former being very much the larger. However, there is no clear geographical
D. J. GRIFFIN AND J. C. YALDWYN 169
dividing line between these two groups and the junction is in the form of a
broad transition zone on the east and west coasts. This is well shown by
plotting the distributions of a large number of tropical and temperate species
on the one map (Fig. 1). Im the case of the majid spider crabs (22 species
occurring mainly in northern waters, 15 in temperate areas), it can be seen
that this transition zone extends on the east coast from about Cape Howe
in the south to Hervey Bay in the north, and on the west coast from about
Cape Leeuwin in the south to about Shark Bay in the north.
About 160 species have a wide northern distribution extending from
north-western Australia to Queensland and sometimes further south to New
g
& Se.
«zd “
eo Ss
eae %°
CS 4e@
& ies
r 5
a i,
te
$ ?
a
=% 3 A
@ a
cf Rs
Fig. 1. Map of Australia showing known distributions of 22 mainly northern
species and 15 mainly southern species of majid crab, indicating the extent of the
transition zones between the northern tropical and southern temperate faunas.
South Wales or Victoria (Tables 3 and 4, Fig. 2). Particularly notable
in this regard are the relatively large number of portunids, leucosiids,
parthenopids, calappids, palinurids, porcellanids and penaeids; a smaller
proportion of the larger families, such as Xanthidae and Majidae, are also
widespread northern forms. It should be pointed out that although the
distributions of these species are considered to be continuous, a very large
number (about 2/3) of these widespread northern species are not known
from the Gulf of Carpentaria and the Torres Strait area.
170 THE AUSTRALIAN DECAPOD CRUSTACEA
More than 40 species have widespread southern distributions extending
from New South Wales or Victoria to south-western Australia. Particularly
notable here are majids, grapsids and hymenosomids; the dromiids are also
well represented in this southern group, although their distributions are more
limited according to the information currently available. Five of these wide-
spread southern species occur from southern Queensland to central Western
Australia.
Less obvious than these northern and southern groups are ones which
are either eastern or western (Fig. 7). The number of species extending
along the whole eastern or western coasts are few and generally eastern
species are either northern or southern. Inasmuch as some species are known
TABLE 3
Number of species showing particular distribution—northern, eastern and southern
Distributional Limits
Famil é : oe i
‘ See ee ee ae 4 ee 8
® dh CA ee 3 pe ae eee
Rate maltose halls oy dlp BOR to <i aie Soe Ul
ere) eerie ne eee Age EO. 7 = = oe
G59 495 9700705 2 Oe
Shae Oram yah - «. Bog S 2 2 ow aa
HS FF 244 eee 4 4 Shree
Xanthidae af 58.0 0b 15.510: 18) 63. =: 2. 2 8 oe a
Majidae we ae 38° 2) TOS De a ee 2 2) ed = OM ee eo
Portunidae .. ae 41 — 12 17 3 2 — 3 — — 2 — — 1 1 —
Leucosidae .. ae 25)... 164 (98 2) oo ers 1—- 2—- 1 ~— 2 —
Paguridae .. . 46 — 2B 2 2 = —. 2 — —. 1 1 eo
Palaemonidae a Wes Th 5 2 1—.— — 2
Grapsidae .. ee 920 cs Ree 2 6 a Se ee SS
Ocypodidae . 22 2 6 5 3 — — 4 1— — —~ ~—~ — — 1
Penaeidae .. i ws 1 2 9 2 — — 1 2 ~—~ — — — J ~—~ —
Porcellanidae ne 12 — 9 2 — — — — 1—-— — —
Alpheidae .. a 8 1 1— —~ 3 —~ —~ —~ —~ 1 ~— — 1 +> 4)
Parthenopidae Be Us Sey me: in (ee 1—- —
Dromiidae .. si 7 — — — — —~ — 1 ~ ~—~ 4 ~~ ~— ~ — 2
Goneplacidae an 4— — — 1 —— 1 — — 1 — > S=S =
Hippolytidae ie 6 — 1 1—- ~—~ ~—~ —~ ~—~ — 8 1 ~—~ ~~ ~— —
Palinuridae .. a 8 — § — ~ —~ ~—~ | ~ ~—~ ~ 1 ~ — 1 —
Calappidae .. a 10 3°05 — — — 1~ ~~ ~ ~ ~—~ 1 ~— —
Others aD fe 40 284 Be BN 8 80 Ae >
Total se 304 18) (87) 162) 51 8 229) 88) 8 2b 1S al See Omni
only from either the east or west coasts there appear to be transition zones
in the north around the Torres Strait area and in the south around eastern
South Australia. The area around Spencers Gulf in South Australia contains
several species which are otherwise common in tropical areas (Fig. 1).
Approximately 130 species have a widespread distribution along the
east coast. The majority of these occur along the Queensland coast often to
northern New South Wales but a large number occur from Queensland or
New South Wales to Victoria, Tasmania or South Australia. One widespread
majid, Oncinopus aranea, extends from northern Queensland to South Aus-
tralia and eight others have only slightly more limited distributions.
Particularly notable in this eastern group are the Xanthidae with a very
large number of species confined to Queensland. At least 20 species of
D. J. GRIFFIN AND J. C. YALDWYN 171
decapods appear to have a central eastern distribution extending from
southern Queensland to northern Victoria.
No separate analysis of western species is given as our distributional
data were summarized under State headings only and subdivision of the
long Western Australian coastline has not been attempted in detail.
_Summarizing the distribution patterns of species occurring Outside a
single State, the most common is a northern one with a smaller group of
species having widespread southern distributions. These two major types are
TABLE 4
Examples of very widely distributed species
Area
Family Species N.
S.A. W.A. Qld. Qld. N.S.W. Vict. S.A. W.A.
Penaeidae .. Penaeus latisulcatus x
Hymenopenaeus sibogae
Palaemonidae .. Anchistus custos x
Alpheidae .. Alpheus edwardsii
Upogebiidae .. Upogebia darwinii x
Hippidae .. Hippa adactyla x
Porcellanidae .. Polyonyx transversus
Pisidia dispar x
Dromiidae .. Petalomera lateralis
Leucosiidae .. Leucosia anatum
Myra mammilaris
Calappidae .. Calappa hepatica
Matuta granulosa
Parthenopidae.. Ceratocarcinus dilatatus
Majidae .. Huenia proteus
Zewa varians
Hyastenus diacanthus
Achaeus brevirostris
Leptomithrax __ sterno-
costulatus ‘
Schizophrys aspera x
5 Micippa philyra x
Xanthidae .. Cryptocoeloma haswelli x
x
x
exex<Sr lee
exaX x
Pex
|
x
x x xX xX X
OOO I ao
Xe Xa lelex
xxxxxxxxXxxXxxXxXX*X
xxx x xX SEX X
x x xX x
x
x
x
x
x
x
x
x
Actaea calculosa
Actumnus setifer
Pilumnus fissifrons
Pilumnus semilanatus x
Ozius truncatus
Portunidae .. Thalamita sima x
Portunus pelagicus x
Macropipus corrugatus
Inssocarcinus polybioides x x =
Grapsidae .. Leptograpsus variegatus
Cyclograpsus audouinit
eo
ll Xeex x
x x
ae
x
x
x
xX X
xX X
XXX XK XKXK XK XK XK
x XXX XK XX
x
Ses oS
eS ON eS
characteristic of particular taxonomic groups. Thus portunids, penaeids,
parthenopids, ocypodids, porcellanids and palinurids are almost entirely
restricted to northern Australia.
A very large number of species in every group is not known from more
than one State. In the case of nine families (Oplophoridae, Crangonidae,
Callianassidae, Paguridae, Majidae, Palicidae, Xanthidae, Goneplacidae and
Grapsidae) more than 2/3 of the species are recorded at present from
only one State.
Different families are represented in quite different proportions in
different areas as can be seen from Tables 3 to 6 and Fig. 3. Figures 4 and 5
give examples of some of the distribution patterns discussed above. The
172 THE AUSTRALIAN DECAPOD CRUSTACEA
families Xanthidae, Majidae and Grapsidae possess several widespread
northern as well as southern species. The Hymenosomidae is the only family
with 10 species or more which is virtually restricted to southern waters
(Queensland species are known, but not, as yet, adequately recorded in the
literature). The different distributions are summarized in Table 3 and Fig. 26.
Extra-AUSTRALIAN GEOGRAPHICAL RELATIONSHIPS
Examination of the distribution patterns of the decapod genera repre-
sented in Australia shows that almost 50% of those found outside Australia
possess species in the Indo-West Pacific area but not outside it, whilst a
very Slightly smaller number are distributed throughout the Indian, Pacific
and Atlantic Oceans. The list below groups the families according to the
relationships of the included genera.
SA WA NGO GO NSw vic SA WA SA WA NGLD GLD NSWw VIC SA WA
————— SN EE ee
SS Se See
SS PRI ETI NE BEN aN PO 8 Tsao ne ee
—
ae 0
oma Rar Cah Si see reate fi
SEE 18
Ee
LRT ED aCe Ge) a b
FP a)
oes a
3 XANTHIDAE a
EEE]
scale - 20 eee ee ere]
50 art
0 —
ae
————= ere es |
ear
is oN)
ae PORTUNIDAE —_——
SR Se eee Lar)
fees even | —s
a aes
[precana tse Nera Sa Se ae nT re]
os EE
Raa) eee Se
a san eR ER
ERS Se eee as
Eason Ss MASIDAE eRe
2 a
ET a
TOTAL MARINE
FAUNA
PENAEIDAE
HYMENOSOMIDAE
Fig. 2. Diagram showing distribution patterns of Australian marine decapod species
known to occur in more than one State; width of each bar is proportional to number
of species showing each particular pattern of distribution according to the scales
given. a—distribution of all Australian species; b—distribution patterns of the species
belonging to five selected families.
Families with more than 2/3 of their Genera World-Wide in Distribution
Large families (i.e., with 10 or more Australian species): Penaeidae,
Scyllaridae, Paguridae, Porcellanidae, Galatheidae, Calappidae.
Small families (i.e., with less than 10 Australian species) : Sergestidae,
Pasiphaeidae, Pandalidae, Oplophoridae, Rhynchocinetidae, Processidae,
Crangonidae, Stenopodidae, Nephropsidae, Axiidae, Upogebiidae, Pylo-
chelidae, Coenobitidae, Chirostylidae, Albuneidae, Homolidae, Dorippidae,
Cancridae, Gecarcinidae, Hapalocarcinidae.
~ D. J. GRIFFIN AND J. C. YALDWYN W763;
Families with more than 2/5 of their Genera Indo-West Pacific in Distribution
Large families (i.e., with 10 or more Australian species) : Leucosiidae,
Majidae, Parthenopidae, Hymenosomidae, Ocypodidae.
Small families (j.e., with less than 10 Australian species): Ogyrididae,
Thalassinidae, Laomediidae, Corystidae, Atelecyclidae, Mictyridae.
Families with between 1/5 and 2/5 of their Genera World-Wide or Indo-West
Pacific in Distribution
. Large families (i.e., with 10 or more Australian species) : Palaemonidae,
Alpheidae, Hippolytidae, Palinuridae, Dromiidae, Xanthidae, Portunidae,
Goneplacidae, Grapsidae, Pinnotheridae.
Small families (i.e., with less than 10 Australian species): Gnatho-
phyllidae, Callianassidae, Hippidae, Raninidae, Palicidae.
TABLE 5
Percentage of total marine decapod species in each state made up by each family
Area—Percentage of Species
Family
Vic. and
Qld. N.S.W. Tas. S.A. W.A. All States
Approx. number of species 700 300 200 200 400 1070
Xanthidae i 166 16 12 8 10 18 15
Majidae sid 93 7 12 18 12 10 9
Portunidae.. 80 10 10 3 4 11 8
Leucosiidae .. 74 8 4 6 5 8 7
Paguridae a 58 4 5 6 2 3 5
Palaemonidae.. . 48 4 2 a 3 3 4
Grapsidae ite 43 5 5 6 4 2 4
Ocypodidae .. 4] 5 3 1 0-5 4 4
Penaeidae m: 51 8 6 + 2 4 4
Porcellanidae .. 38 a 2 3 1 7 3
Alpheidae Sif 31 3 2 4 7 1 2
Parthenopidae. . 29 4 1 _ 1 2 2
Dromiidae aie 24 1 2 3 6 2 2
Goneplacidae .. 21 2 3 3 2 0-5 1-5
Hippolytidae .. 19 0-5 1 1-5 3 1-5 1:5
Scyllaridae ae 15 1 2 1 — 1-5 1-5
Calappidae sm 14 5 2 — 1-5 3 1-5
Pinnotheridae 12 1 0-5 1-5 1-5 1 1
Palinuridae .. 11 1 1-5 1 0-5 1-5 H
Galatheidae .. 11 1 2 2 0-5 0-5 I
Hymenosomidae 10 — 1 2-5 2-5 1 1
Others vA 111 13 21 24-5 34 13-5 21
Families and Genera with Asiatic Representation
The families Atyidae and Potamidae (both freshwater), and representa-
tives of the following families: Majidae (Leptomithraxz, Sargassocarcinus) ;
Xanthidae (Hypothatessia, Calvactea, Liagore) ; Grapsidae (Utica) ; Palicidae
(Pleurophrycus).
Genera with a Wide, Southern Temperate Distribution
Genera, with family name in brackets, as follows: Nawticaris (Hippo-
lytidae) ; Campylonotus (Campylonotidae) ; Jasus (Palinuridae) ; Noto-
mithrax (Majidae); Halicarcinus (Hymenosomidae); Leptograpsus and
Cyclograpsus (in part) (Grapsidae) ; Nectocarcinus (Portunidae).
174 THE AUSTRALIAN DECAPOD CRUSTACEA
Some Genera with Other Distributions
Indopacific and South Atlantic: Muwrsia (Calappidae).
Indopacific and West African: Hpixanthus, Heteropanope, Pilumnopeus
(Xanthidae).
Indopacific and West Indies: Chlorodiella, Carpilius, Phymodius
(Xanthidae) ; Ovalipes (Portunidae).
Indopacific and Eastern Pacific (but not Atlantic): Pugettia (Majidae) ;
Daira, Carpilodes, Trapezia (Xanthidae).
More than two thirds of the included genera in almost one half of the
families occur throughout the world, whilst 1/5 of the families are mainly
Indo-West Pacific. Slightly less than one third of the families have mixed
relationships. A few genera, each including only a small number of species
TABLE 6
Number of species per state eapressed as a percentage of the total number of species in each family
Approx. Area—Percentage of Species
Number
Family of Vic. and
Species Qid. N.S.W. Tas. S.A. W.A.
Xanthidae SE os 166 61 20 7 10 42
Majidae Bi: aye ae 93 48 36 34 23 4]
Portunidae .. fe Me 80 75 38 10 7 56
Leucosiidae .. Ss Ae 74 71 15 13 12 46
Paguridae .. es ss 58 40 26 15 U 19
Palaemonidae eo ee 48 48 Il 0 11 22
Grapsidae .. is de 43 80 38 22 20 21
Ocypodidae Ze aps 4] 81 25 5 2 47
Penaeidae .. ae a 51 - 90 35 11 9 34
Porcellanidae ic = 38 61 16 7 5 74
Alpheidae .. Bie Se 31 60 16 20 Ai 20
Parthenopidae os fe 29 80 10 0 10 29
Dromiidae .. a i 24 31 26 21 40 31
Goneplacidae Ae a 21 49 40 20 20 5
Hippolytidae Bi v6 19 15 15 15 40 30
Scyllaridae .. Ld ae 15 51 48 12 0 34
Calappidae .. ake ie 14 70 50 t) 22 80
Pinnotheridae ae ie 12 40 9 25 25 45
Palinuridae .. bi a ll 85 61 24 10 75
Galatheidae i a 11 85 72 48 9 18
Hymenosomidae .. im 10 0 40 50 50 40
Others dis es Ke Ill 90 50 70 65 60
in Australia, are distributed mainly through the southern temperate areas of
the Indo-Pacific. The freshwater families, Atyidae and Potamidae, as well
as a few marine genera, are largely confined to Australia and the Indo-
Malayan area. Finally, a small number of genera, mainly xanthids, have
unusual distributions.
Twenty-nine genera are not known outside Australia; many of these are
monotypic. The following is a list of these restricted and often poorly known
taxa.
Genera Endemic (i.e., restricted) to Australia
Freshwater genera and families: Caridinides, Stygiocaris (Atyidae) ; ten
genera of Parastacidae (i.e., all except Cheraz).
Marine genera, with family names in brackets, as follows: Vercoia
(Crangonidae); Lomis (Lomisidae); Epipedromia (Dromiidae); Lisso-
Sl
D. J. GRIFFIN AND J. C. YALDWYN 17
PERCENT OF TOTAL SPECIES PER STATE
Os © pO © 1/0» Of-OrcCoy o' ©
‘> UITYVTNANUTT is _|
1O.0.0 0.02620 -010 100-0. 000-6, x +
qeeectetetet ete tetatetetetetet = > Oo
rg re
Ga =
= =| >
g aot tS
m O
>
S$ mi
Y SOOO IOOOORORO 8]
OOO OOO OOOO
Vererereceretetetecee wee
o
Oo Fv
D> Om
= >
=> m
— 7 BIUUIITTNIIIIINILIIV NULL 5 Oo
U Se
m
: m
Mm 1O-0.0.6.0,0'6.0,0.0-0'0,
Srorerererereretere rere.
SSS S85
S |
PX KDODODOODDOD
OOOO POOXOOE™
SSSI
D0020000
RSS 525052
Fig. 3. Histogram showing percentage of total species per family per State for
each of four selected families compared with percentage State representation of total
Australian marine decapod fauna.
THE AUSTRALIAN DECAPOD CRUSTACEA
b
Fig. 4. a—Distribution of the stridulating group of penaeid prawns of the genus
Metapenaeopsis showing various patterns of northern distribution. b—Distribution of
of the ocypodid crab Hemiplax latifrons, the majid crab Naxia aurita and the grapsid
crab Leptograpsus variegatus showing three patterns of southern distribution.
a a
D. J. GRIFFIN AND J. C. YALDWYN ALT/
“Il
Fig. 5. a—Distribution of the palinurid crayfish Panulirus cygnus and the majid
crab Paranaxia serpulifera showing two types of western distribution pattern; and of
the callianassid “yabby” Callianassa australiensis and the scyllarid lobster Arctides
antipodarum showing two types of eastern distribution pattern. b—Distribution of the
raninid crab Lyreidus tridentatus and the majid crab Schizophrys aspera showing two
types of discontinuous distribution.
178 THE AUSTRALIAN DECAPOD CRUSTACEA
morpha (Leucosiidae) ; Tumulosternum, Paramithraz, Paranaxia, Ephippias
(Majidae) ; Leptograpsodes, Helograpsus, Paragrapsus (Grapsidae) ; Banarei-
opsis, Paraetisis, Pseudocarcinus, Prolybia (Xanthidae) ; Heloecius, Austral-
oplax (Ocypodidae).
TROPICAL AND TEMPERATE SPECIES AND THEIR DISTRIBUTIONS
An analysis of the distributions of the species belonging to five family
groups (Table 7) show that a large proportion of the tropical species are
widespread throughout the Indo-West Pacific. Generally a smaller proportion
is either shared with part of the Indian Ocean or the Western Pacific, though a
sizeable number of Australian species are not known outside the Indo-Malayan
Archipelago—in the penaeine prawns this last group is particularly large.
The temperate group is characterized by a very large number of species
which are restricted to Australia; among the temperate majids a small
percentage is shared with southern temperate areas outside Australia.
TABLE 7
Distributions of species outside Australia for five family groups
Percentage of Total Species per Distribution
Type Partitioned according to Area
1)
E N= = ©
Family 5 5 4 3 5 : ap
Gas aro eres ee a et se ee
= 3; ' Ba 15 g a lyme)
oS) ~ e) “1 © = ~ Oy
29 zg cs ae o = 3 5
o 8 ° I aS iS iS) Dy,
Sa «8 sO ff m Mo
ie v4 19 10 ll 8 52 0 91 . Tropical Australia
Bereunidaensen 180. 66 0 0 0 34 0 9 Temperate Australia
Natid. ‘ 17 15 10 15 43 0 65 Tropical Australia
MEMES ae 67 0 2 11 6 14 35 Temperate Austraha
Oxe stomata 45 17 4 24 9 46 0 82 Tropical Western Austraha
ahs grrr 66 6 22 0 2 0 18 Temperate Western Aust-
ralia
Porcellanidaeso 21 11 12 18 38 0 88 Tropical Western Austraha
100 0 0 0 0 0 12. Temperate Western Aust-
raha
PenAGnae 40 38 2) 8 15 18 0 92 Tropical Australia
ie e, 100 0 0 0 0 0 8 Temperate Austraha
The fauna as a whole is not as yet well enough known to allow an analysis
of the representation of the main Indo-Pacific elements in the various areas
of Australia. Thus in the majids, whilst at present the western Australian
fauna appears to contain a larger percentage of Indian Ocean species than
does the eastern Australian fauna, this may be due to the fact that the
western Australian fauna is the poorer known.
CONCLUSIONS—PRESENT AND FuturRE WorK
Much of what has been outlined above would apply to most groups of
Australian marine animals—tropical groups tend to be widely distributed,
whereas cool temperate animals tend to have limited distributions.
Zoogeographically, the Australian decapod fauna appears to have been devived
almost entirely from the tropical Indo-Pacific region. There seems to be no
z D. J. GRIFFIN AND J. C. YALDWYN 179
good evidence to suggest that temperate species have reached Australia
directly from temperate areas outside Australia.
The following concluding remarks attempt to summarize what is at
present known about each of the main systematic groups. It should be
pointed out that few parts of the Australian coastline, continental shelf
and off-shore oceanic waters have been well sampled thus far, whilst the
continental slope beyond 100 fms. is virtually unknown. Almost the whole
of the area around north-western Australia and the Great Australian Bight
is unexplored.
_The penaeid prawns are continuing to be actively investigated by
A. A. Racek of the University of Sydney. Almost nothing is known about
the identity and distribution of the pelagic and bathypelagic shrimps
(Sergestidae, Pasiphaeidae, Oplophoridae), nor about the shelf and archi-
benthal, possibly commercial, natants (e.g., Pandalidae and Crangonidae) ;
only the Challenger Expedition (1873-76) has provided as yet any substantial
information on these groups in Australian offshore waters. However, New
Zealand representatives have been recently reviewed (Richardson and
Yaldwyn, 1958; Yaldwyn, 1960).
Among the most complex and taxonomically difficult groups of natants
are the Alpheidae (snapping shrimps—see Banner and Banner, 1966), the
Australian members of which are currently being revised by A. H. and Dora
M. Banner of the University of Hawaii. The other main natant group—the
Palaemonidae—is perhaps reasonably well known; the pontonines (mainly
tropical commensals—see Bruce, 1967) are now being studied by A. J. Bruce
of the C.S.1.R.O. Division of Fisheries and Oceanography and _ the
palaemonines have been briefly investigated by a few workers (e.g., Holthuis,
1952). Our comprehension of the freshwater species of the genus Macro-
brachium and of the freshwater shrimps of the family Atyidae, is almost
abysmal—nothing definitive is known of the individual distributions, or of
the range of variation in these forms. The general zoogeography and relation-
ships of all freshwater decapod genera has, however, been reviewed in detail
by Bishop: (1967).
For most of the reptant macrurans our knowledge is more advanced. The
status of the palinurids is probably better known than that of any other
group of decapods, and they are under continuing study by R. W. George
of the Western Australian Museum and L. B. Holthuis of the Rijksmuseum,
Leiden (George and Holthuis, 1965). Holthuis has also nearly completed
revisionary studies on the ScyHaridae. Two major systematic revisions of the
Parastacidae are being completed by E. F. Riek of the ©.8.1.R.O. Division
of Entomology and D. D. Francois of the N.S.W. Chief Secretary’s Depart-
ment Fisheries Division.
Not a great deal is known of the very large and varied group, the
Anomura. Only the Porcellanidae are reasonably well known, the western
species having recently been studied by Janet Haig of the Allan Hancock
Foundation, Los Angeles (Haig, 1965) and she is soon to commence intensive
studies on the remainder of the Australian fauna. Information about the
pagurids is almost totally absent for the whole Indo-Pacific (with the excep-
tion of south-east Asia where the species have been studied by Fize and
Seréne, 1955); this is in contrast to the fairly well-known Atlantic fauna
(see Gordan, 1956). The position of the monotypic Lomisidae has been
investigated recently by Pilgrim (1965).
In the Brachyura a few groups have been studied in varied detail, but
most are still poorly known. Knowledge of the leucosiids, calappids and
180 THE AUSTRALIAN DECAPOD CRUSTACEA
raninids has been given a basis in the recent studies by Marina Tyndale—
Biscoe and R. W. George (1962), and by W. Stephenson of the University
of Queensland (pers. comm.). These oxystomes are one of the groups where
further revisionary studies could be completed fairly easily in the near future.
In the case of the oxyrhynchs (Majidae, Parthenopidae and Hymen-
osomidae), one of us (D.J.G.G.) is continuing investigation of the majids;
a fairly large number of new species remain to be added to the fauna and
many species known from eastern Australia are present in western waters,
but have not so far been recorded formally. Studies on the parthenopids have
been initiated; the most recent relevant studies are by Flipse (1930) and
Serene (1954, 1955). The Hymenosomidae exhibit their greatest diversity in
Australasia (Tesch, 1918@), yet knowledge of them is almost entirely lacking;
the work of J. 8S. Lucas, of the University of Western Australia (pers. comm.),
on reproductive isolation and general biology, should provide a sound basis
for future work on the Australian species.
The portunids continue to undergo intensive study by W. Stephenson
and his co-workers at the University of Queensland; this group is one of the
best known of Australian crabs. The very large and complex group, the
Xanthidae, are currently being revised on a world-wide basis by Daniéle
suinot of the National Museum, Paris (see Forest and Guinot, 1961) ;
previous knowledge of Australian species comes mainly from the works of
Melbourne Ward (see Whitley, 1967). The Goneplacidae appear at present
to make up rather a small proportion of the brachyuran fauna but it should
be pointed out that several small and taxonomically difficult subfamilies
await study; many Indo-West Pacific species have been recently documented
by Seréne (1964).
The behaviourally and ecologically very interesting shore crabs of the
families Grapsidae and Ocypodidae are perhaps reasonably well known in
the Indo-Pacific (Banerjee, 1960; Crosnier, 1965) and several groups are
currently under review in Australia. Local species of the first family are
being studied by B. M. Campbell of the Queensland Museum (Campbell,
1967; Campbell and Griffin, 1966) who is particularly interested in the
complex group belonging to the genus Sesarma. R. Seréne of the National
Museum, Singapore, is revising at the same time the Indo-Pacific species of
Sesarma, and one of us (D.J.G.G.) is investigating the distribution of, and
variation in, several widespread shore crab species (Griffin, in press). The
macrophthalmine ocypodids are currently being revised by R. S. K. Barnes
(1966a, 6). The fiddler crabs (Uca species) are in an unusual position
taxonomically (see Crosnier, 1965); a very sound basis for work on these
species was provided by studies on American species in the 1940’s by Jocelyn
Crane who has more recently briefly investigated most of the Indo-Pacific
species (Crane, 1957). Several workers in Australia are variously interested
in this group and know something of the distribution and status of local
species, based mainly on recent ecological studies by W. Macnae (1966) of
Witwatersrand University, Johannesburg. However, nothing substantial is
at present being done by anybody on Australian members of this genus. The
western Australian ghost crabs (Ocypode species) have recently been studied
by R. W. George and Mary Knott (1965).
Nothing is known of two interesting families of mainly commensal species;
the Pinnotheridae (see Tesch, 1918); Seréne, 1964), which undoubtedly con-
tain many more Australian species than are currently recognised and the
Hapalocarcinidae of which almost nothing is known in Australia, although
D. J. GRIFFIN AND J. C. YALDWYN 181
the south-east Asian fauna has recently been thoroughly documented by Fize
and Seréne (1957).
Undoubtedly a lot of work remains to be done. Most importantly,
knowledge must be obtained of the distributions and patterns of variation
within species. Work along these lines might be considered as preceding
systematic rearrangements and generic revisions in those cases where identi-
fication of taxa at the generic level is not impossible. It is to be earnestly
hoped that currently available knowledge concerning some groups will be
fully exploited in the near future in conjunction with ecological studies.
Species exist in nature, not merely in a bottle or in a person’s mind, it is
the systematist’s job to find them.
References
BAtss, H., 1935.—Brachyura of the Hamburg Museum expedition to south-western Aus-
tralia, 1905. J. r. Soc. W. Aust., 21: 113-151, 5 figs, pl. 13.
, 1957.—Systematik. In Bronns, H. G., Klassen und Ordnungen das Tierreichs
5 (1) 7, Decapoda, 12: 1505-1672, figs 1131-1199.
BANERJEE, S. K., 1960.—Biological results of the Snellius expedition. XVIII. The genera
Grapsus, Geograpsus and Metopograpsus (Crustacea Brachyura). Temminckia,
10: 132-199, 6 figs.
Banner, A. H., and Banner, Dora M., 1966.—Contributions to the knowledge of the
alpheid shrimp of the Pacific Ocean—Part X. Collections from Fiji, Tonga and
Samoa. Pacif. Sci., 20: 145-188, 20 figs.
Barnes, R. S. K., 1966a.—A new species cof the genus Macrophthalmus Latreille, 1829
(Decapoda: Brachyura:Ocypodidae) from the Gulf of Carpentaria, Queensland.
Proc. r. Soc. Qd, 78: 438-48, 1 fig., pl. VIII.
, 19666.—The status of the crab genus Huplax H. Milne Edwards, 1852; and a
new genus Australoplax of the subfamily Macrophthalminae Dana, 1851 (Brachyura:
Ocypodidae). Aust. Zool., 13: 370-376, pl. XXIV.
BisHop, J. A., 1963.—The Australian freshwater crabs of the family Potamonidae
(Crustacea: Decapoda). Aust. J. mar. freshwat. Res., 14: 218-288, 4 figs, 2 pls.
, 1967—The zoogeography of the Australian freshwater decapod Crustacea.
In WEATHERLEY, A. H., “Australian Inland Waters and their Fauna.” Australian
National University Press, Canberra. Pp. 107-122, 6 figs.
Bruce, A. J., 1967—Notes on some Indo-Pacific Pontoniinae III-IX. Descriptions of
some new genera and species from the western Indian Ocean and the South China
Sea. Zool. Verhand., 87: 1-73, 29 figs.
CAMPBELL, B. M., 1967.—The Australian Sesarminae (Crustacea: Brachyura). Five species
of Sesarma (Chiromantes). Mem. Qd Mus., 15: 1-19, 2 figs, 5 pls.
, and Grirrin, D. J. G. 1966—The Australian Sesarminae (Crustacea:
Brachyura): genera Helice, Helograpsus nov., Cyclograpsus and Paragrapsus. Mem.
Qd Mus., 14: 127-174, 10 figs, pls XX-X XIII
CLARK, Hllen, 1936.—The freshwater and land crayfishes of Australia. Mem. natn. Mus.
Vict., 10: 5-58, 11 pls.
= 1941.—Revision of the genus Huastacus (Crayfishes, Family Parastacidae),
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182 THE AUSTRALIAN DECAPOD CRUSTACEA
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THE EMBRYOLOGY OF EPALTES AUSTRALIS LESS.
(COMPOSITAE)
Gwenpa L. Davis
Department of Botany, Uniwersity of New England, Armidale, N.S.W.
[Read 24th April, 1968]
Synopsis
The capitula of H. australis are heterogamous and, although all the florets are
tubular, those of the outermost seven or eight whorls are filiform and female. Along
any radius of the capitulum, the anthers develop in acropetal succession whereas the
ovules form and mature in a basipetal manner.
Microsporogenesis is regular and the pollen is three-celled when shed.
Embryo sac formation follows the Polygonum type but only two antipodal cells are
formed and the micropylar cell is binucleate. No secondary multiplication of antipodals
occurs and the cells persist into post-fertilization stages. Fusion of the polar nuclei
occurs immediately before fertilization and after the premature degeneration of the
synergids.
Endosperm formation is ab initio Cellular and embryogeny conforms to the Asterad
type.
INTRODUCTION
Of the three species of Hpaltes endemic to Australia, EH. australis is the
most widely distributed and, according to Black (1957), it occurs in all states
except Western Australia. It has also been reported from Formosa where it
was possibly introduced in Australian wheat or wool.
The plant is herbaceous with procumbent stems which may reach 30 cm.
in length and the hemispherical capitula are borne on short pedicels in the
axils of the obovate leaves.
No member of this genus has previously been the subject of an embryo-
logical investigation although its component species are endemic to tropical
Africa, Mexico, Brazil, tropical Asia and China, as well as Australia.
MATERIALS AND MertHops
The material on which this investigation was based was collected from
40 miles east of Wanaaring, far western New South Wales, and fixed in
F.A.A. by Professor N. C. W. Beadle of the University of New England.
After embedding, sections were cut at 8-10u and stained with Delafield’s
haematoxylin and Johansen’s safranin.
FLoraLu MorpeHouocy
The capitula are hemispherical and heterogamous and the naked
receptacle is slightly concave at maturity. The florets are all tubular and
do not exceed the ovate, obtuse, involucral bracts, their number varying with
the size of the capitulum. According to Black (1957), about 100 female
florets surround the 8—25 bisexual disc florets but, in the Wanaaring material,
PROCEEDINGS OF THE LINNEAN Society oF New SoutH WALES, Vout. 93, Part 1
GWENDA L. DAVIS 185
the ratio of female to bisexual florets was approximately 12:1 in a total of
up to 400 florets per capitulum. .
The filiform female florets occupy the outermost 7 or 8 whorls and, at
maturity, the stylar arms protrude through the small aperture at the apex
of the corolla tube (Text-fig. 1). Their outer surfaces are finely papillate and
no definite stigmatic lines could be identified at their margins (Text-fig. 3).
The bisexual disc florets are shortly 4-lobed (Text-fig. 2) and the filaments:
of the four epipetalous stamens are inserted near the base of the corolla
tube. Although the anthers are closely associated, they do not cohere into
an anther tube. The stylar arms are lanceolate and taper sharply into the
slender apices (Text-fig. 4). Finely papillate stigmatic lines occupy their
margins and the large papillae on their outer surfaces are continuous with
those on the distal end of the style proper. A ring-like nectary surrounds
the base of the style and has no counterpart in the female florets.
The fruits of the bisexual florets are slightly broader than those of the
females but both types are longitudinally ribbed and bear a microscopic rim
which represents the pappus (Text-figs 1-2).
MiIcrROSPORANGIUM
The anthers are tetrasporangiate with sterile apices and are proximally
tailed. Within the epidermis the fully formed anther wall is made up of
endothecium, middle layer and tapetum and its method of formation follows
the Dicotyledonous type (Davis, 1966). The cells of the amoeboid tapetum
become multinucleate during meiosis in the adjacent microspore mother cells,
and this may be accompanied by nuclear fusion. When the microspore tetrads
are formed, the tapetal cells are 2- or 4-nucleate and, on breakdown of the
tetrads, periplasmodium formation occurs.
Microsporogenesis and male gametogenesis follow the same sequence of
events as described in Podolepis jaceoides (Davis, 1961) and the pollen grains
are 3-celled when shed after the longitudinal dehiscence of the anther.
MEGASPORNAGIUM
The initially atropous ovule becomes anatropous with the development
of the single massive integument which overgrows the nucellus until it meets
the funicle (Text-figs 5-7). As in other Compositae, the ovule is tenuinucellar
and arises from the base of the inferior ovary.
Megasporogenesis
An archesporial cell differentiates at the apex of the nucellus in a sub-
epidermal position and gives rise directly to the megaspore mother cell
(Text-figs 5-7). The enlargement of this cell is accompanied by active cyto-
plasmic synthesis and it is accommodated in the nucellar lobe by anticlinal
divisions of the overlying nucellar epidermal cells.
When the growth of the integument is completed, meiosis is initiated in
the megaspore mother cell whose nucleus is situated towards the micropylar
pole (Text-figs 8-9). Cytokinesis after meiosis 1 is therefore unequal! and the
chalazal dyad cell is larger than its micropylar counterpart (Text-fig. 10).
Meiosis 11 takes place simultaneously in both dyad cells (Text-fig. 11) and
is followed by wall formation but, whereas the two micropylar megaspores
are the same size, the chalazal dyad cell divides unequally and the chalazal
megaspore of the linear tetrad is the largest (Text-fig. 12). Although all the
THE EMBRYOLOGY OF EPALTES AUSTRALIS LESS
186
development of the embryo sac. Mm, micro-
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GWENDA L. DAVIS 187
Mmegaspores may develop vacuoles and, consequently, are said to germinate,
only the chalazal megaspore increases in size and the three non-functional
megaspores degenerate (Text-fig. 13).
Embryo sac formation
‘When the functional megaspore enlarges, the nucellar epidermis becomes
tangentially stretched and its cells do not divide anticlinally. The transition
from the functional megaspore stage to the 1-nucleate embryo sac is purely
a matter of size increase and is not associated with any anatomical charac-
teristics. In most Compositae, the 1-nucleate embryo sac fills the nucellar
lobe after the dissolution of the non-functional megaspores, but in H. australis
the latter are still in evidence. when the nucleus of the embryo sac divides
(Text-fig. 14). A vacuole forms between the daughter nuclei and, following
the eventual absorption of the crushed non-functional megaspores, the
2-nucleate embryo sac comes into direct contact throughout its length with
the nucellar epidermis (Text-fig. 15). Further enlargement of the embryo
sac results in its apex penetrating the nucellar epidermis and taking up its
position at the throat of the micropyle (Text-fig. 16). Although the cell
contents of the nucellar epidermis are in a degenerated condition, the layer
itself persists until maturity of the embryo sac and its remains can be
recognised internal to the endothelium.
The second embryo sac mitosis leads to the 4-nucleate stage in which
the nuclei occupy the 2+2 configuration, the two pairs being separated by
a large vacuole which was formed after the preceding nuclear division (Text-
figs 17-18)..The third and last embryo sac mitosis, as well as the following
free 8-nucleate stage, were not observed but, after cytokinesis, differentiation
occurs and the development of the embryo sac is, therefore, of the Polygonum
type.
The differentiated embryo sac is 6-celled (Text-fig. 19), due to the forma-
tion of only two antipodal cells, of which the micropylar is invariably
2-nucleate. In the newly-formed embryo sac, the antipodal cells are vacuolate
and their cytoplasm stains lightly but, at maturity, the vacuoles disappear
and each cell becomes filled with dense cytoplasm. No nuclear divisions occur
and the two cells remain in this condition until they degenerate when embryo-
geny is well advanced. In the endosperm mother cell, the two polar nuclei
initially occupy opposite poles and are separated by the large vacuole which
has persisted since the 2-nucleate stage of the embryo sac. The chalazal polar
nucleus then migrates to the micropylar portion of the cell where it becomes
closely associated with its micropylar counterpart. Fusion of the polar nuclei
to form a secondary nucleus is delayed until just before fertilization. As in
other Compositae, the three cells of the egg apparatus are partially enclosed
by the hood-like micropylar portion of the endosperm mother cell. When the
synergids are first formed, they are slender basally vacuolate cells which
taper towards their apices at the throat of the micropyle. However, during
their degeneration, each develops a lateral fold which appears hook-like in
section (Text-fig. 20). In Hpaltes australis the breakdown of the synergids
is premature in that it commences before the formation of the secondary
nucleus and is not associated with the entrance of the pollen tube into the
embryo sac. The egg is an elongated cell which is deeply enfolded by the
endosperm mother cell and its micropylar half is occupied by a large vacuole
(Text-figs 20-21).
The micropylar chamber, which contains the nucellar lobe, is bounded
by the innermost cell layer of both the integument and the raphe. These cells
AUSTRALIS LESS
THE EMBRYOLOGY OF EPALTES
188
Text-figs 19-24—Mature embryo sac, fertilization and endosperm formation. e, egg;
m, male gametes; pt, pollen tube in micropyle; s, sygergids; v, vegetative nucleus;
z, zygote. Text-figs 19 x 600; 20, 21, 23, 24 x 433; 22 x
67; 22a, 22b x 1000.
= GWENDA L. DAVIS 189
divide only anticlinally (Text-figs 10, 12) and, by the time the megaspore
mother cell enters meiosis, they have formed the endothelium. In cell
arrangement and staining capacity, this cell layer is sharply demarcated
from the remainder of the integumentary cells and it reaches its maximum
development at maturity of the embryo sac. Its cells show signs of degenera-
tion after endosperm formation has been initiated and it can be distinguished
only by its position by the time the young embryo has become heart-shaped.
The development of an endothelium is invariable in the Compositae and it is
a character of tenuinucellar ovules in general, where the embryo sac comes
into direct contact with the integument after the breakdown of the nucellus.
Circumstantial evidence indicates that it may play a part in the nutrition
of the embryo sac by producing enzymes which digest the contents of the
integumentary cells external to it. In text-figure 22 this process of digestion
is well advanced and a region of empty collapsed cells extends outwards
from the endothelium, within which is the mature embryo sac.
Fertilization
Syngamy was not observed, but in one ovule (Text-fig. 22) in which a
secondary nucleus was present and the synergids had degenerated (Text-fig.
22a), a pollen tube was present in the micropyle and contained two closely
associated vermiform male gametes as well as the vegetative nucleus from
the pollen grain (Text-fig. 22b).
POST-FERTILIZATION EVENTS
Endosperm formation
Division of the primary endosperm nucleus precedes that of the zygote.
The mitotic spindle lies at right angles to the long axis of the embryo sac
and a wall is laid down across the telophase spindle (Text-fig. 28). Endosperm
formation is therefore of the ab initio Cellular type and subsequent divisions
follow rapidly. By the time the proembryo is 2-celled, endosperm formation
is well advanced (Text-fig. 24) but there is no evidence of its digestion before
the embryo has reached the stage shown in text-figure 35. When the embryo
is fully grown all the endosperm has been utilized with the exception of a
layer of regular thick-walled cells within the remains of the integument. The
seed, therefore, is not strictly exalbuminous.
Embryogeny
After formation of five or six endosperm cells, the zygote divides trans-
versely to form the superposed cells ca and cb of the two-celled proembryo
(Text-fig. 25). Vertical division of ca gives rise to the tier gq and cb divides
transversely to form the cells m and ci of the four-celled proembryo (Text-
figs 26-27). A vertical division in q and m converts the first into a quadrant
and the second into two juxtaposed cells, while ct divides transversely to
form n and n’ (Text-figs 28-29). The eight-celled proembryo is now made up
of four tiers of cells, g, m, nm and n’; further vertical divisions transform m
into a quadrant and replace the single cell of m by two juxtaposed cells
(Text-figs 30-31). This is the final stage of the proembryo and, after trans-
verse division of n’ into 0 and p, the four-tiered embryo proper is delimited
from the suspensor which originates from p. Because ca undergoes a vertical
division and cells derived from both ca and cb take part in the formation
of the embryo proper, the embryogeny of H#. australis conforms to the Asterad
190 THE EMBRYOLOGY OF EPALTES AUSTRALIS LESS
type. The quadrant q now becomes an octant and a periclinal division of
each cell of m cuts off a central quadrant (Text-fig. 32). The cells of q divide
periclinally, further vertical and periclinal divisions occur in m, n becomes.
a quadrant and p forms p’ and p” (Text-fig. 33). Transverse divisions cause
m to become two-tiered and the three-celled suspensor (ss) is established
(Text-figs 34-35). The embryo proper has now assumed a globular form and
Aiea G
saaies
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RE
se
Text-figs 25-38—Embryogeny. (Lettering follows the system of Souége.) Text-figs
PASI 6 (INS Oko xe CRBS Biri 3¢ PINS aX) b< GAO), s
the suspensor cells become highly vacuolate and of irregular outline. Their
contents disappear and, although the suspensor can still be identified when
the embryo becomes heart-shaped (Text-fig. 36), no trace of it remains after
the initiation of the cotyledons (Text-figs 37-38).
MATURATION GRADIENTS IN THE CAPITULUM
The capitulum being a racemose inflorescence, its florets mature in acro-
petal succession and the youngest occupy a central position. There is therefore
a gradient expressed in the maturation of both anthers and ovules along any
radius. In H. australis, however, although the stages of microsporogenesis
were found to conform to this centripetal pattern, a reverse gradient was
observed in the development of the ovules. In all the capitula examined the
central florets contained the youngest anthers and the most mature ovules,
= GWENDA L. DAVIS 191
whereas the peripheral female florets contained the youngest ovules. This
phenomenon of opposite maturation gradients in the male and female repro-
ductive structures of a capitulum has not been reported previously and, in
order to investigate it more closely, the ovules of florets occupying three
sites in median sections of five capitula were compared:
Central Bisexual Innermost Female Outermost Female
Floret Floret Floret
1. Growth of integument almost Integument development Ovule atropous
completed initiated
2. Megaspore tetrad ‘Meiosis 11 Meiosis 1
3. 2-Nucleate embryo sac Functional megaspore Megaspore tetrad
4, 4-Nucleate embryo sac _ First embryo sac mitosis Functional megaspore
5. 2-Cellerd poembryo 2-Celled proembryo Mature embryo sac
Due to their earlier maturation, the florets at the centre of the capitulum
are the first to be pollinated and proembryos are found in their ovules before
fertilization occurs in the peripheral florets. During embryogeny, the matura-
tion gradient becomes less apparent and, in the infructescence, the ovaries of
all florets contain a seed with a fully developed embryo.
DISCUSSION
Although embryo sac formation follows the Polygonum type, cytokinesis
is such that invariably the chalazal antipodal nucleus is enclosed in one cell
and the remaining two in the second. This nuclear disposition is in agree-
ment with all other reports in the Compositae where there are only two anti-
podal cells and there is no record of the chalazal cell ever being the binucleate
one. Variation does occur, however, in the behaviour of these cells. For
example, in Synedrella nodiflora (Banerji and Pal, 1955) and Tridax trilobata
(Hjelmqvist, 1951) fusion may occur between the nuclei in the micropylar
cell so that both cells are then uninucleate, and in Bidens biternata
(Deshpande, 19646) and Gerbera jamesonii (Maheswari Devi, 1957) the nuclei
divide and each cell becomes multinucleate. The commonest condition, how-
ever, is where nuclear division is accompanied by cell division. This secondary
multiplication of antipodal cells has been reported by Harling (1951) in
Chrysanthemum arcticum, Erigeron canadensis and Matricaria globifera and
later (1954) in Vittadinia triloba. In Epaltes australis no embryo sac was
observed with more than two antipodal cells and, although the two nuclei
in the micropylar cell were closely associated, fusion did not occur. In this
respect, this species is similar to Chrysanthemum flosculum (Harling, 1951),
Flaveria repanda (Misra, 1957), Matricaria chamomilla (Harling, 1951),
Senecio glutinosus (Afzelius, 1924), Tridaxw procumbens (Maheshwari and
Roy, 1952), Volutarella ramosa (Deshpande, 1964a) and Wedelia calendulacea
(Ghosh, 1962) although the occasional occurrence of three antipodal cells is
reported in these examples.
The premature degeneration of the synergids is an unusual feature of the
embryology of EL. australis. This appears to be an autonymous breakdown of
the cells because it occurs before the entry of the pollen tube and while the
embryo sac is still immature. No comparable examples of this phenomenon
have been traced in the literature.
192 THE EMBRYOLOGY OF HPALTES AUSTRALIS LESS
References
AFZELIUS, K., 1924.—Hmbryologische und zytologische Studien in Senecio und ver-
wandten Gattungen. Acta Hort. Berg., 8: 123-219.
BAnergi, I., and SunANDA Pat, 1959.—A contribution to the life history of Synedrella
nodiflora. J. Linn. Soc. Lond. (Bot.), 55: 810-817.
Biack, J. M., 1957.—Flora of South Australia. 2nd Ed. (Govt. Printer, Adelaide).
Davis, G. L., 1961.—The life history of Podolepis jaceoides. 1. Microsporogenesis and
male gametogenesis. Phytomorphology, 11: 86-97.
, 1966.—“‘Systematic Embryology of the Angiosperms.” (John Wiley, New York.)
DESHPANDE, P. K., 1964a.—A contribution to the life history of Volutarella ramosa.
J. Indian bot. Soc., 43: 141-148.
, 1964b.—A contribution to the embryology of Bidens biternata. J. Indian bot.
Soc., 48: 149-157.
GHOoSH, R. B., 1962.—A contribution to the life history of Wedelia calendulacea. J. Indian
bot. Soc., 41: 197-206.
HARLING, G., 1951.—Embryological studies in the Compositae. 1-3. Acta Hort. Berg.,
16: 1-120.
, 1954.—The embryo sac development of Vittadinia triloba. Svensk bot. Tidsk.,
48: 489-496.
HsrectmMevist, H., 1951.—The embryo sac development of Tridax triloba. Bot. Notiser,
1951: 180-187. :
MAHESHWARI, P., and Roy, S. K., 1952.—The embryo sac and embryo of Tridax pro-
cumbens. Phytomorphology, 2: 245-252.
MaAneESswaArRt Devi, H., 1957.—Embryological studies in the Compositae. 3. Gerbera
jamesonit. Proc. Indian Acad. Sci. B., 46: 68-74. :
Misra, 8., 1957.—Floral morphology of the Compositae. 1. The flower and gametophytes
of Flaveria repanda. J. Indian Bot. Soc., 36:503-512.
AUSTRALASIAN MEDICAL PUBLISHING CO. LTD. —
71-79 ARUNDEL ST., GLEBE, SYDNEY, N.S.W., 2037
APHROPHYLLUM (RUGOSA) FROM LOWER CARBONIFEROUS
LIMESTONES NEAR BINGARA, NEW SOUTH WALES
R. K. JuLu* ,
University College of Townsville, Townsville, Queensland
(Communicated by Mr. R. H. Anderson)
(Plate x1I1)
[Read 26th June, 1968]
Synopsis
The hystero-ontogeny of the type species of Aphrophyllum, A. hallense Smith, is
described and A. smithi, sp. nov. is proposed. Both species are known only from
Viséan (Lower Carboniferous) limestones at Halls Creek, near Bingara. Some similari-
ties between the youthful and adult characters of A. hallense and those of Thysano-
phyllum orientale Nicholson and Thomson, type species of Thysanophyllum, and
Lonsdaleia spp. suggest that Aphrophyllum may lie with the Lonsdaleiidae; other
features, however, suggest that it lies in a separate, new family of rugose corals.
INTRODUCTION
Studies of Lower Carboniferous corals from New South Wales by Pickett
(1967) and work in progress by the writer on Queensland forms have shown
Aphrophyllum Smith to be an important and diverse genus endemic to the
Viséan of eastern Australia. Its affinities are in doubt, with Smith (1920)
and Jones (1933) suggesting a possible relationship with Hndophyllum
Edwards and Haime, Hill (1956) and Dobrolyubova (1962) relating it to
Palaeosmilia Edwards and Haime, and Pickett (1967) placing the genus with
the Lonsdaleiidae. With the intention of clarifying this problem, the follow-
ing discussion is devoted to the details of the hystero-ontogeny of its type
species A. hallense Smith. Aphrophyllum smithi, sp. nov., which is also
described herein, provides some measure of the variability of the genus in
the area of provenance of its type species.
Material for this study is from limestones at Halls Creek, some 17 miles
south of Bingara, New South Wales (Text-fig. 1). Pickett (1967) considered
these beds to be Middle Viséan in age and correlated them with the upper
part of the Namoi Formation.
All fossil and locality numbers (“F” and “L” numbers) are registered
in the catalogue of the Department of Geology and Mineralology, University
of Queensland, Brisbane with the exception of those prefixed with “AM”
Which are in the Australian Museum, Sydney, and with “BM”, in the British
Museum (Natural History), London. The terminology applied to hystero-
ontogeny is that as outlined by Jull (1965).
SYSTEMATIC DESCRIPTION
Genus APHROPHYLLUM Smith
APHROPHYLLUM HALLENSE Smith
(Pl. x11, figs 1-3; Text-fig. 2)
Holotype: AM F17640 (not F17648 or F17684 as quoted respectively by
Hill, 1934, p. 73 and Pickett, 1967, p. 29) with thin sections AM1036-38-39-40
in the Australian Museum, Sydney (listed as B& by Smith, 1920, p. 64)
and A5051 in the Sedgwick Museum, Cambridge.
* Now at the Department of Geology, University of Windsor, Windsor, Ontario,
Canada.
PROCEEDINGS OF THE LINNEAN Society or NEw SoutH WALES, VoL. 93, Part 2
194. APHROPHYLLUM (RUGOSA) FROM LOWER CARBONIFEROUS LIMESTONES
Locality and Age: The species is known only from a few localities at
probably nearly the same stratigraphic horizon in Viséan limestones in the
vicinity of Halls Creek, in Parish Hall, County Murchison, some 17 miles
south of Bingara, New South Wales. Smith (1920, p. 51) in proposing
the species, did not list a specific locality in these limestones for the holotoype
and paratype. However, Benson who collected the material, reported (1917,
p. 242) a fauna in this limestone in Portion 46, Parish Hall. As this is his
only mention of a fossiliferous limestone in the area, presumably this is the
type locality. I did not visit this spot during my trip to the area in 1964,
but Mr. F. W. Mitchell, property owner on Halls Creek, who provided accurate
information on other outcrops of the limestone, described to me the occurrence
of a fairly prominent limestone outcrop immediately west of Red Rocks, a
locality marked in Portion 46 of the Parish Map (see Text-fig. 1). This is
possibly coincident with Professor Benson’s collecting site.
i Graftone
eBingara
®8Map Area x L2817
e Tamworth
\ /
\ Portion 46, |
1 »)
\Parish of Hally
Newcastle
Scale in miles
Bingara Falls_>
Text.-fig. 1. Map of the Halls Creek area showing localities of specimens collected.
Site of the type locality of Aphrophyllum hallense Smith is probably near “Red Rocks”.
Material for this study was collected from two localities in this lime-
stone. One (2816) is 300 yards north along strike from Bingara Falls and
about three-quarters of a mile south of the presumed type locality. The other
(L2817) is on the east side of Halls Creek, 25 yards west of the serpentine
Ie, 1S AS OILIO, 195
and approximately one and one-half miles north of Benson’s site. The lime-
stones outcrop discontinuously in the area but the three localities are likely
to prove to be nearly at the same horizon. There is no doubt that all material
in question is conspecific.
Diagnosis: Cerioid, with irregularly shaped corallites averaging, 18 mm.
in diameter; usually 20 to 24 septa of each of two orders are present; the
long often pinnately arranged major septa sometimes meet at the axis to
form an irregular axial structure, and the minor septa are well developed;
lonsdaleoid dissepiments occupy a variably wide zone and are naotically
modified in part; tabulae are domed with flat or upturned ends; increase is
lateral, rarely parricidal peripheral; trabeculae are monacanthne.
Description: A. Adult characters. Smith (1920), Hill (1934) and
Pickett (1967) have all described the adult characters in adequate detail.
Pickett (1967) placed in Nothaphrophyllum gregarium Pickett the specimen
from Chatham Quarry, Taree, considered by Hill (1934) to belong to the
present species.
B. Hystero-ontogeny. All known representatives of A. hallense are more
or less crushed and this, together with the abundant stereome in the species
which tends: to obscure detail, and the fact that the axial septa are not
obvious, makes the study of corallite development difficult. Amongst eleven
coralla, only 46028 from L2816 yielded nearly complete details of hystero-
ontogeny from examination of closely spaced acetate peel sections of the
specimen. The following description is based principally on one corallite of
this specimen, illustrated in Text-fig. 2. In general detail, this example of
corallite development is typical of other youthful corallites which were
observed in thin sections from other specimens including the holotype. Figure
references, viz. F46028/18—28, refer to details in Text-fig. 2.
Lateral increase.
Increase is typically lateral but parricidal peripheral increase has been
observed in one instance, this being described after the section on lateral
increase. The laterally arising corallite developed within the lonsdaleoid
dissepimentarium of the parent and all of its septa are independently inserted ;
they do not represent continuation of growth of parts of the septa of the
parent. Stereome is abundant throughout development.
Hystero-brephic stage (F46028/18—23). The daughter corallite is first
obvious as a small area within the lonsdaleoid dissepiments of the parent,
and approximately 0:25 mm. distally axial septa appear. The axial plane
is orientated radially to the axis of the parent with the cardinal septum
on the outer side, furthest from the axis of the parent. Axial septa are
initially united but almost immediately separate into counter and cardinal
septa which never come into direct contact with each other again during
development. Simultaneous with their separation is the appearance of. alar
septa, followed shortly by counter-lateral septa. After the shortening of the
axial septa, none of the six septa are in contact during this stage of develop-
ment which ranges over approximately 0-45 mm. of growth.
Hystero-neanic stage (F46028/24-38): This stage commences with the
appearance of metasepta, and includes most of the septal insertion in the
daughter corallite. Insertion is somewhat irregular, with septa arising from
_ the wall of the corallite in fossulae which are not obvious. Newly inserted
Septa are not contratingent.
Most major septa are inserted before minor septa in a regular manner
in the right cardinal quadrant. Insertion in the left cardinal quadrant would
196 APHROPHYLLUM (RUGOSA) FROM LOWER CARBONIFEROUS LIMESTONES
probably have been similar had not this side of the corallite been constricted
during development with the result that major septum number 4 in this
quadrant is late in appearing.
Insertion in the counter quadrants is characterized by periods of rapid
insertion of major and minor septa. In general, major septa alternate with
minor septa in order of appearance. A disproportionately large number of
septa are inserted in the left counter quadrant (the reverse of the diagram-
matic corallite in Text-fig. 2), resulting in the lateral migration of the counter
CARDINAL QUADRANTS foECTION NO. F46028/— COUNTER QUADRANTS
eS at \
3
114
\
25 6n 713
| | |
25
mm.
|
|
|
|
|
|
|
3
3
Text-fig. 2. Hystero-ontogeny in Aphrophyllum hallense Smith, F46028, from L2816.
The diagrammatic representations of cardinal and counter quadrants illustrate the
order of septal insertion; note the change in vertical scale. C= cardinal septum;
K= counter septum; A= alar septum; solid lines= long major septa; short dashed lines=
short major septa; long dashed lines= minor septa. Number 1 septa in the cardinal
quadrants are alar septa. Numbers opposite the diagrammatic quadrants are those
of pertinent acetate peel sections. Numbers 18 to 23 are through the hystero-brephic
stage, 24-38 through the hystero-neanic stage, 39-72 through the late neanic stage, and
73 through the ephebic stage. Lower five corallites « 2:2, upper three corallites x 1:1.
septum during corallite growth so that axial septa cease to lie opposite each
other. Possibly it was adjacent to the left counter quadrant that the daughter
met with the least resistance to corallite expansion.
During the early part of the hystero-neanic stage, major septa do not
reach the axis, but they lengthen with further corallite development so that
Rae UE 197
by the end of the hystero-neanic stage, some extend to the axis and an
irregular axial structure is consistently present.
Throughout the hystero-neanic stage, the daughter corallite is subcircular
in outline and lonsdaleoid dissepiments are absent, except for the occasional
small one. Interseptal dissepiments are absent at first, but by the end of
this stage of development, a single row is developed entirely around the
corallite.
Approximately 2°75 mm. of growth occurs during the hystero-neanic
stage. The end of this stage and the commencement of the late neanic stage
are transitional.
Late neanic stage (F46028/39-72) : The daughter corallite now acquires
sharp corners and some lonsdaleoid dissepiments. A few pairs of major septa
alternating with minor septa in order of appearance, are inserted at the wall
in the cardinal and alar fossulae. A small axial structure of twisted lamellae
is present until just before the achievement of the ephebic stage; in some
corallites this axial structure persists into adulthood. Little development of
the lonsdaleoid dissepimentarium occurs until the later part of this stage.
At this time, a wide zone of lonsdaleoid dissepiments develops around the
corallite, and many of these dissepiments become naotically modified. With
this development, the corallite reaches adulthood, after 20 mm. of growth
during this stage and a total of about 23 mm. of growth.
Parricidal peripheral increase.
In the observed example cf parricidal peripheral increase, four corallites
arose in the calice of the parent, and these soon expanded to occupy the
entire area of the calice. A fifth corallite appeared somewhat later than the
rest and may have arisen by lateral increase from its developing neighbour.
The early stages of development of these corallites have been destroyed by
crushing. In general, their hystero-ontogeny resembles that observed in the
above described example of lateral increase, except that a lonsdaleoid dis-
Sepimentarium is developed earlier, but nevertheless not widened nor
naotically modified until the final stage of development. Three of these
corallites are illustrated in the lower-left of pl. x1, fig. 1.
Remarks: The variable nature of the adult morphology of the species is
likewise reflected in the nature of corallite development, and it is difficult
to assess which features of the latter are typical of the species. Certainly
the character of axial septa ceasing to oppose one another after early develop-
ment in the example illustrated is one of individual variation, and so also is
the exact pattern of septal insertion. However, the rather general pattern
during hystero-neanic development of major septa appearing before the minor
septa in the cardinal quadrants, and major and minor septa appearing alter-
nately in the counter quadrants, is probably fairly typical of the species; it
somewhat resembles insertion in laterally arising corallites of Lithostrotion
(see Jull, 1965). Other apparently significant characters are the initially -
united axial septa which are radially orientated with respect to the axis of
the parent during hystero-brephic development; and the late neanic appear-
ance of an irregular axial structure and of lonsdaleoid dissepiments which
are mainly formed as such and not as modifications from interseptal
dissepiments.
Increase in A. hallense differs in a number of characters from that in the
type species of Thysanophyllum, T. orientale Thomson and in Lonsdaleia spp.
(see Jull, 1967). United axial septa during hystero-brephic development, and
minor septa appearing in the counter quadrants at the start of the hystero-
198 APHROPHYLLUM (RUGOSA) FROM LOWER CARBONIFEROUS LIMESTONES
neanic stage are unknown in 7’. orientale and Lonsdaleia. They also generally
lack the formation of lonsdaleoid dissepiments per se, these structures
forming rather from interseptal dissepiments by the withdrawal of septa
from the wall.
In lateral increase in Lonsdaleia, commonly as many as five corallites
develop nearly simultaneously from the same parent and some of these coral-
lites are orientated with their axial planes as much as 90 degrees away from
the parent; neither of these characters have yet been observed in either A.
hallense or T. orientale.
APHROPHYLLUM SMITHI, Sp. Nov.
(Pl. x11, figs 4-5)
Holotype: F46073, from L2816, located in Viséan limestones at Halls
Creek 300 yards north along strike from Bingara Falls, Portion 62, Parish
Hall, County Murchison, New South Wales.
Diagnosis: Similar to A. hallense but with smaller corallites having fewer
septa, a narrower lonsdaleoid dissepimentarium, and flatter tabulae.
Description: Based on the holotype and three topotypes, all of which
have been partially crushed.
The corallum is cerioid and ponsists of irregularly shaped polygonal
corallites which average 11 mm. and range from 7:5 to 14 mm. in diameter.
Intercorallite walls are sinuous or undulating as seen in transverse section
of the corallite, and are 0-4 to 0-5 mm. in width. Stereome lines the dis-
sepiments and in some corallites, it may almost totally infill the interstices
of the skeletal elements.
Septa are straight or slightly wavy and have irregular outlines. Corallites
have from 18 to 22 septa of each order. Major septa generally do not reach
the axis in the holotype, but in most corallites of the other specimens (pl. x1,
fig. 5) they extend close to the axis or reach it. Long septa are pinnately
arranged in elongate corallites. An axial structure is absent. Minor septa
are less than half as long as major septa.
The dissepimentarium consists of one to three rows of steeply inclined
intersepta! dissepiments bounded by variably developed lonsdaleoid dissepi-
ments which are usually crushed. Lonsdaleoid dissepiments are developed
entirely around the corallite, or are present only in a narrow zone in the
corners or ends of corallites. Naotic developments are uncommon. The
tabularium ranges from 5-5 to 7 mm. in diameter. Tabulae are gently arched
or domed if septa extend close to the axis, or flat or sagging if septa are
short. The ends of tabulae are flat or uptur ned, and approximately 20 tabulae
occur in a 10 mm. interval.
Hystero-ontogeny: The few youthful corallites observed appear to have
arisen by lateral increase. They are similar to those in A. hallense, but an
axial structure was not observed.
Microstructures: Septal trabeculae are monacanthine and similar to those
in A. hallense (see Text-fig. 4) except that trabeculae are more closely spaced,
being at approximately 0:075 mm. intervals, rather than 0-1 to 0-125 mm. as
they are in A. hallense.
Distribution: The species is known only at the type locality.
Remarks: The species is named in honour of Dr. Stanley Smith who first
described corals from the limestones at Hails Creek. Apart from possessing
lip 1X5 AOAC, 199
. SMITHI
Te 9 0 nm D 13 14°15 16 17 18 49 20 2 2 23
56
54
52
50
- 48 12
ts elie dial aie
NS. 12 ae
44 6 ee 4
® A 12
4 Ga
G Tegal ale |
40 6 /® 15 A. SMITHI ®
38 \6 4—B5 b—
\ 15 16
36 \ @ P,
4
3 2 44 3B 38 4D ad wSSOBSOSO SD GA <S6 686062 G4 GS GB 7
T cp,
56 56
54 : 54 T | ke
52 52 9
50 ph 50 7 + |
4 1 3 48 3)\aeen
46 1210 46 + +—12 10 _
A. HALLENSE| |
Av. 44 Salle 8 7 = Av. 44 ies A.HALLENSE © |78
N.S. | SY | NS. A ~ % |
42 ® |+—_———_—_—_+ !—_}+— @>
N
40 A.SMITHI
38
36
34
30
56 58 60 62 64 66 68 70 72 7A 76 78 80 92
Av. T.0.
Cc D
Text-fig. 3. Graphical comparison of Aphrophyllum smithi, sp. nov. and A. hallense
Smith. N.S. = number of septa of both orders; C.D. = corallite diameter (the mean
of the maximum and minimum dimensions in mm.); T.D. = tabularium diameter
(in mm.); Av. = average. Hach corallum studied was assigned a number, and all
corallites measured in a corallum are recorded on the graphs with the number of their
particular corallum. Vertical numbers are of corallites in A. hallense, and inclined
numbers are of A. smithi. Circled numbers are of corallites in the holotypes. Only
uncrushed or slightly crushed corallites were measured.
200 APHROPHYLLUM (RUGOSA) FROM LOWER CARBONIFEROUS LIMESTONES
smaller dimensions overall, A. smithi is similar to A. hallense, and larger
than average corallites in this species may be mistaken for A. hallense
although corallites of the latter usually have a wider lonsdaleoid dissepimen-
tarium and more strongly arched tabulae. The two are graphically compared
in Text-fig. 3, and some overlap in parameters is evident, especially in graph B.
AFFINITIES OF APHROPHYLLUM
Although the adult and youthful morphology of A. hallense is now well
known, the affinities of the genus are still not readily evident. This is partly
due to the paucity of information on the hystero-ontogeny of rugose corals
in general so that the significance of particular characters is yet not fully
appreciated.
Aphrophyllum was considered by both Hill (1956, p. F290) and
Dobrolyubova (1962, p. 316) to be related to Palaeosmilia; Hill placed the
genus in the Amygdalophyllinae.
For the following reasons, however, the affinities of the genus might
appear to be with the Lonsdaleiidae, as was first suggested by Pickett (1967).
The tabulae of A. hallense and A. smithi resemble those in Thysanophyllum
orientale; this last species, the type of Thysanophyllum, was concluded by
Jull (1967) to lie in the Lonsdaleiidae. When naotic developments are absent
in Aphrophyllum, a common situation in this genus, the lonsdaleoid dissepi-
mentarium is similar to that in typical lonsdaleid corals. Moreover, corallites
of Aphrophyilum in which septa do not reach the axial region closely resemble
the typical morphology of Thysanophyllum; this especially applies to some
undescribed Queensland species, such as Aphrophylluwm sp. illustrated by Hill
and Woods (1964, pl. C2, fig. 1).
Other characters of Aphrophyllum, on the other hand, are unknown in
typical representatives of Lonsdaleia and Thysanophyllum. These are the
pinnate arrangement of septa in all known species of Aphrophyllum, some
details of the hystero-ontogeny of A. hallense, as discussed above, and the
nature of the axial structure.
Septa in Aphrophyllum, Lonsdaleia and Thysanophyllum are composed
of monacanthine trabeculae (Text-fig. 4). In A. hallense, trabeculae are larger
than is normal in these other genera, and the peripheral ends of septa in
some corallites are composed of two or more rows of trabeculae and may
be split or cavernous. Multiple rows of trabeculae are a common situation
in some corals showing naotic developments, but are unknown in Lonsdaleia
and Thysanophyllum. Wang (1950) has earlier remarked on septal structure
in these genera.
There is thus a strong suggestion that Aphrophyllum lies in a separate,
new family of rugose corals. Further work on other species of Aphrophyllum
is currently in progress by the writer and may assist in elucidating the
taxonomic position of this genus.
Finally, Smith (1920, p. 55) and Jones (1933, p. 60) have both remarked
on the similarity of Aphrophyllum hallense to Endophyllum; superficially at
least, the former differs significantly only by having naotically modified dis-
sepiments. Hndophyllum is widely distributed in Middle to Upper Devonian
beds and three cerioid species described by Gorsky (1935) from the
Tournaisian of Novaya Zemlya appear to belong to the genus (see Soshkina
and Dobrolyubova, 1962). Possibly hystero-ontogenetic studies of Hndo-
phyllum may show that Aphrophyllum is descended from this line.
Re Rey i Ua 201
Y Acknowledgements
I am very grateful to Professor D. Hill, F.R.S., both for her criticism
of the manuscript of this paper and her advice during my studies at the
University of Queensland, of which this paper is part of the outcome. Mr.
O. H. Fletcher, formerly of the Australian Museum, Sydney, helpfully pro-
vided photographs and information on the type material of A. hallense, and
I gladly acknowledge the assistance of Dr. B. Runnegar, of the University
of New England, during our visit to the Halls Creek area in 1964 to collect
the present material.
Text-fig. 4. Monacanthine septal trabeculae in A, B, Aphrophyllum hallense Smith,
F46043 from 12816, and in C, D, Lonsdaleia floriformis floriformis (Martin), B.M.
R17160, from Viséan D, zone beds at Coalbrookdale, Shropshire, England. All figs
approx. x 7.
References
Benson, W. N., 1917.—The geology and petrology of the Great Serpentine Belt of New
South Wales. Pt. 4. A general account of the geology and physiography of the
western slopes of New England. Proc. Linn. Soc. N.S.W., 42: 223-283, pls 18-20.
DoprotyuBova, T. A., 1962.—Suborder Caniniina; In ‘‘Principles of Palaeontology” (U. A.
Orlov, Ed.). Spongia, Archaeocyatha, Coelenterata, Vermes (B. S. Sokolov, Ed.),
314-317. Acad. Sci. U.S.S.R. (in Russian).
202 APHROPHYLLUM (RUGOSA) FROM LOWER CARBONIFEROUS LIMESTONES
Gorsky, I. I., 1935.——Some Coelenterata from Lower Carboniferous deposits in Novaya
Zemlya. Trans. arctic Inst., 28: 1-128, pl. 1-12 (in Russian with English summary).
Hitt, D., 1934—The Lower Carboniferous corals of Australia. Proc. roy Soc. Qd.,
45: 63-115, pl. 7-11.
, 1956.—Rugosa; In “Treatise on Invertebrate Palaeontology” (R. C. Moore, Ed.).
Part F, Coelenterata, 233-324. Univ. Kansas Press and Geol. Soc. Amer.
, and Woops, J. T., 1964.—Carboniferous fossils of Queensland. Qd. Palaeontogr.
Soc., 32 pp., 15 pls.
Jones, O. A., 1933.—A revision of the Australian species of the coral genera Spongo-
phyllum BH. & H. and Endophyllum EH. & H. with a note on Aphrophyllum Smith.
Proc. roy. Soc. Qd., 44 (for 1932): 50-63, pl. 3-4.
JULL, R. K., 1965.—Corallum increase in Lithostrotion. Palaeontology, 8: 204-225.
, 1967.—The hystero-ontogeny of Lonsdaleia McCoy and Thysanophyllum
orientale Thomson. J[bid., 10: 617-628, pl. 100-102.
Pickett, J., 1967.Lower Carboniferous coral faunas from the New England district
of New South Wales. Mem. geol. Surv. N.S.W., Palaeont., 15: 38 pp., 20 pls.
SMITH, S., 1920.—On Aphrophyllum. hallense gen. et sp. nov. and Lithostrotion from the
neighbourhood of Bingara, N.S.W. J. Proc. roy Soc. N.S.W., 54: 51-65, pl. 2-5.
SosHKina, H. D., and Doprotyupova, T. A., 1962—Order Columnariida;In “Principles
of Palaeontology” (U. A. Orlov, Ed.). Spongia, Archaeocyatha, Coelenterata,
Vermes (B.S. Sokolov, Hd.), 339-344. Acad. Sci. U.S.S.R. (in Russian).
Wane, H. C., 1950.—A revision of the Zoantharia Rugosa in the light of their minute
skeletal structures. Phil. Trans. roy. Soc. (Ser. B), 234: 175-246, pl. 4-9.
EXPLANATION OF PLATE XIII
All figures x 2.
Figs 1-3. Aphrophyllum hallense Smith. 1, transverse section of F46028 from
L2816; 2, transverse section of topotype, AM 1037; 3a-b, transverse and longitudinal
sections of F46071 from L2817.
Figs 4-5. Aphrophyllum smithi, sp. nov. 4a—b, transverse and longitudinal sections
of holotype, F46073 from L2816; 5, transverse section of topotype F46074 from L2816.
Proc. Linn. Soc. N.S.W., Vol. 98, Part 2 PLATE X1iI1
Aphrophyllum hallense Smith and A. smithi, sp. nov.
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TYPE SPECIMENS IN THE MACLEAY MUSEUM,
UNIVERSITY OF SYDNEY
I. FisHes
P. J. STANBURY
The Macleay Museum, School of Biological Sciences, University of Sydney
(Communicated by Dr. D. T. Anderson)
[Read 26th June, 1968]
INTRODUCTION
The Macleay collection was started around 1790 by Alexander Macleay
(1767-1848). He collected insects. He caught specimens himself and acquired
others by exchange among his friends; but probably the major part of his
collection resulted from purchases at auctions of historic collections. In this
way Macleay obtained specimens originally collected by Captain Cook’s
naturalists, Surgeon General White, Charles Sturt and Sir Stamford Raffles as
well as many other less well known figures (Fletcher, 1920).
Alexander Macleay brought his collection to Australia in 1825-26 when
he was appointed the first Colonial Secretary. His son, William Sharp
Macleay (1792-1865), helped his father collect and after William Sharp
inherited it, he enlarged it considerably. When he died the collection passed
to his cousin, Sir William Macleay (1820-1891), the first President of the
Linnean Society of New South Wales.
Sir William Macleay decided to expand the collection to include all
branches of the Animal Kingdom as well as anthropological and geological
specimens. He employed collectors all over the world and amassed a huge
collection that overflowed from his large house at Elizabeth Bay, Sydney,
into a museum-building 115 x 36 feet in the garden. When this became full,
and having no heir, he decided to offer the entire collection to the University.
of Sydney. The Senate accepted Sir William’s gift in 1888 and the Govern-
ment contributed £16,000 to erect a Museum building in the University
grounds. The collection was moved in 1889, and the building was opened to
the public in 1890.
Sir William Macleay died in 1891. The curator he had appointed, George
Masters, died in 1912. In 1917, the University started to use part of the
building for other purposes. The collection became neglected and its space
was severely curtailed as described by my predecessor (Anderson, 1965). In
the last few years, however, some attention has been given to the Museum
and the following table gives an idea of what remains.
An account of the history of the Macleays was given by J. J. Fletcher
in his Presidential Address to the Linnean Society of New South Wales,
14th June, 1920. This account was completed after Fletcher’s death from
his notes by A. B. Walkom (Fletcher, 1929). The story of Sir William
Macleay’s collecting trips has been told by D. N. 8S. MacMillan in his book
“A Squatter Went “ae Sea” (1957).
PROCEEDINGS OF THH LINNEAN Society or NEw Souru WaAtEs, Vor. 93, Part 2
204 TYPE SPECIMENS IN THE MACLEAY MUSEUM
So far as I can discover, the only lists of type specimens in the collection
that have been published are:
(i) CopLanp, S. J., 1945, 1946.—Catalogue of Reptiles in the Macleay
Museum, Parts 1 and 2. Proc. Linn. Soc. N.S.W., 70: 291-811; 71:
136-144. These papers dealt in detail with three species of lizards.
(ii) Haun, D. E.—A list of the designated type specimens in the Macleay
Museum: Insecta. Duplicated at the University of Sydney, 1962.
This listed about 2,000 type specimens.
TABLE 1
An estimate of the number of specimens in the Macleay Museum,
University of Sydney, in January, 1968
Approximate
Specimen Total Number of
Types
Coelenterates .. oe 300 a=
Sponges .. aH be 200 3
Corals 7s fe 45 250 =
Annelids .. ce Sr 100 =
Insects .. aif ae 1,000,000 2,000 +
Shells ¥: aS si 50,000 150
Other molJluses .. ie 300 ae
Echinoderms ps ais 500 =
Fishes us oe bie ~ 9,000 75
Amphibians is ae 1,000 7
Reptiles .. a oe 2,000 60
Birds ts as ic 9,000 12+
Mammals ae of 1,500 8
Anthropological “fs 2,000 Many
Fossils we 200 Some
I propose to publish lists of the other type specimens in the Macleay
Museum. This will be done in the following order: Fishes; Amphibians
Reptiles; Birds; Mammals; and Shells.
This paper contains a list of the fishes of the Macleay Museum from
which new species were described.
THe Fisu Types
Nearly all the fish types in the Museum were described by Sir William
Macleay. The original descriptions were published in the Proceedings of
the Linnean Society of New South Wales. Most of them were reprinted at
Sydney in 1881 by F. W. White, in the “Descriptive Catalogue of Australian
Fishes” (Vols. I and II).
The bulk of Sir William Macleay’s type specimens are in the Macleay
Museum, although about 40 are in the Australian Museum, Sydney (Whitley,
1957).
Macleay collected fishes from specially chartered vessels such as the
“Galatea” and the “Peahen” which dredged up and down the coast of New
South Wales (Fletcher, 1929), and the “Chevert? which sailed into New
Guinea waters (MacMillan, 1957). Macleay also acquired unusual fish from
the Sydney Fish Market and from friends.
The following list includes all the fishes that I can find in the Museum
that Macleay described as new species. Most of his names have since been
P. J. STANBURY 205
superseded, some many times. Further types may lie unrecognised in the
Macleay Museum. The fish are stored in bottles in 70% alcohol.
The classification in the list is based on that given by D. S. Jordan,
“The Genera of Fishes and a Classification of Fishes” (1963) and the family
numbers are Jordan’s. The F numbers are the Macleay Museum Register
numbers.
The data for each type in the list that follows is arranged in the following order:
Family name/Family number/Scientific name under which described/Kind of type/
Number of specimens/Locality/Reference in Proc. Linn. Soc., N.S.W./Macleay Museum
register number.
Hemiscyllidae 51 Chiloscyllium furvum Macleay H 1 Port Jackson VI. 364, No 1094,
1881 F7
' Squalidae 67 Acanthias megalops Macleay H 1 Port Jackson VI 367, No 1101, 1881 F24
Myliobatidae 84 Myliobatus australis Macleay H 1 Port Jackson VI. 380, No 1124, 1881
F41(a)
Dussumieriidae 148 Etrumeus FOES OniCneis Macleay C 2 Port Jackson III. 36, 1878 F66
Galaxiidae 173 Galazias bongbong Macleay C 10 Moss Vale VI. 233, No 849, 1881 F82
Galaxiidae 173 Galazrias nebulosa Macleay C 2 Long Bay nr. Sydney VI. 234, No 850,
1881 F83
Galaxiidae 173 Galaxias planiceps Macleay C 9 Rankin’s Lagoon; Bathurst VI. 233,
No 848, 1881 F85
Galaxiidae 173 Galaxias coxiti Macleay C Approx. 14 Mount Wilson VI. 232, No 847,
1881 +V p. 46, 1880 F87
Sternoptychidae 185 Sternoptychides amabilis Ogilby C Damaged H 1 +bits Lord Howe
Island III. pt. 3, p. 1313, 1888 F93
Alabidae 197 Chilobranchus rufus Macleay C Approx. 15 Port Jackson VI. 266, No 909
1881 F96
Alabidae 197 Chilobranchus rufus Macleay C Approx. 15 Tasmania VI. 266, No 909
1881 F96(a)
Echelidae 216 Muraenichthys australis Macleay C 2 Lane Cove VI. 272, No 921, 1881
F114
Echelidae 216 Myrophis chrysogaster Macleay H 1 Port Darwin VI. 271, No 918, 1881
F115
Echelidae 216 Myropterura laticaudata Ogilby C 2 Fiji XXII. 247, 1879 F116
Plotosidae 245 Copidoglanis longifilis Macleay C 2 Long Is. Torrest Straits. “Chevert”
Exp. 1875 VI. 207, No 809, 1881 F158
Synodontidae 271 Saurida truculenta Macleay H 1 Port Jackson VI. 219, No 830, 1881
F174
Synodontidae 271 Saurida argentea Macleay C 2 Endeavour River VI. 220, No 830, 1881
F175
Belonidae 302 Belone gracilis Macleay C 4 Port Jackson VI. 243,- No 865, 1881 F190 °
Gadidae 310 Lotella marginata Macleay C 2 Port Jackson VI. 114, No 767, 1881 F209
Bothidae 326 Rhomboidichthyls spiniceps Macleay H 1 Port Jackson VI. 127, No 783,
1881 F211
Bothidae 326 Arnoglossus bleekeri Macleay H 1 Endeavour River VII. 124, No 779,
1881 F217
Bothidae 326 Lophorhombus cristatus Macleay H 1 Port Jackson VII. 14, 1882 F1160
Paralichthyidae 327 Teratorhombus excisiceps Macleay H 1 Port Jackson VI. 126, No 782,
1881 F218
Soleidae 333 Solea macleayana Ramsay C(?) 1 Port Jackson V. 462, 1880 F1167
Synapturidae 334 Synaptura sclerolepis Macleay H 1 Port Darwin IT. p. 363, pl. 10, fig. 4,
1877 F225
Synapturidae 334 Synaptura nigra Macleay C 3 Port Jackson V. 1881 48-9 F225 (a)
Synapturidae 334 Synaptura fasciata Macleay H 1 Port Jackson VII. 1882 14 F1168
Cynoglossidae 335 Plagusia guttata Macleay H 1 Endeavour River II. 362, pl. 10, fig a
1877 F226
Cynoglossidae 335 Plagusia guttata Macleay C 6 Port Darwin II. 362, pl. 10, fig 3,
1877 F226(a)
Cynoglossidae 335 Plagusia unicolor Macleay C 2 Port Jackson VI. 138, No 801, 1881
F1162
Holocentridae 348 Holocentrum goldiei Macleay C 3 Port Moresby VII. p. 352, No 127,
1882 F241
’
)
206 TYPE SPECIMENS IN THE MACLEAY MUSEUM
Syngnathidae 356 Leptoichthys cristatus Macleay H 1 Western Australia VI. 296,
No. 964, 1881 F256(a)
Syngnathidae 356 Ichthyocampus maculatus Alleyne and Macleay H 1 Darnley Is.
“Chevert” Exp. 1875 I. 3538 pl. 17, fig 2, 1875 F261
Syngnathidae 356 Ichthyocampus annulatus Macleay C 2 Port Darwin II. 364, pl. 10,
fig 6, 1875 F262
Synenathidae 356 Stigmatophora depressiuscula Macleay H 1 King George’s Sound
VI. 299, No 969, 1881 F280
Centriscidae 362 Amphisile komis Macleay H 1 Yap Is. III. 1876 p. 165 F290
Melanotaeniidae 372 Aristeus rufescens Macleay C 2 Rivers of Northern Queensland
V. 625, No 5388 1880 F294(a)
Melanotaeniidae 372 Aristeus lineatus Macleay H 1 Richmond River V. 626, No 539,
1880 F294(b)
Melanotaeniidae 372 Atherinosoma jamesonii Macleay C 6 Bremer R. Qld. IX. 171, 1884
F295
Mugilidae 374 Mugil delicatus All and Macleay C 4 Cape York “Chevert” Exp. 1875
I. 341, pl. 15, fig. 1, 1875 F306
Polynemidae 376 Polynemus caecus Macleay H 1 Port Darwin II. 354, pl. 9, fig. 1,
1877 F325
Thunnidae 378 Pelamys australis Macleay H 1 Port Jackson V. 557, No 397, 1880
F333
Carangidae 401 Caranx edentulus All and Macleay C 4 Percy Is. I. 327, pl. 11, fig. 2 1875
F349
Carangidae 401 Caranx bucculentus All and Macleay C. 2 Cape York I. 326, pl. 11, fig. 1,
1875 F350
Carangidae 401 Caranx radiatus Macleay H(?)° 1 Rockingham Bay V. 537, No. 362,
1880 F351
Carangidae 401 Caranz cheverti All and Macleay H 1 Katow; New Guinea, 1875 I. 324,
pl. 10, fig. 1, 1875 F357
Carangidae 401 Caranx obtusiceps eee C 9 Port Moresby VII. 357, No 151, 1882
F360
Carangidae 401 Caranz mandibularis Macleay C 2 Port Moresby VII. 356, No 150, 1882
F363
Carangidae 401 Caranz laticaudis All and Macleay H 1 Hall Sound, New Guinea 1875
1.8325, pl. 10; fis: 2:°1875 W365
Carangidae 401 Caranxz papuensis All and Macleay H 2 Hall Sound, New Guinea I. 325,
pl. 10, fies 3s 1875 W366 ;
Carangidae 401 Caranaz moresbiensis Macleay H 1 Port Moresby VII. 358, No 152, 1882
F369
Apogonidae 414 Apogon guttulatus All and Macleay C Approx 20 Darnley Island.
“Chevert” Exp. 1875 I. 267, pl. 5, fig. 1, 1875 F402
Apogonidae 414 Apogon cookii Macleay C 10 Endeavour River V. 344, No 88, 1880 F403
Apogonidae 414 Apogon cookii Macleay C(?) 7 Port Moresby V. 344, No 88, 1880 F409
Apogonidae 414 Apogon opercularis Macleay C 12 Port Darwin II. 347, pl. 7, fig. 1,
1877 F406
Apogonidae 414 Apogonichthys darnleyensis All and Macleay H 1 Darnley Is. ‘“Chevert”
Exp. 1875 I. 268, pl. 5, fig. 3, 1875 F418
Apogonidae 414 Apogonichthys marmoratus All and Macleay C. 2. Cape Grenville
T3685) (ple be. tices llSii5 eral
Apogonidae 414 Apogonichthys roseobrunneus Macleay H 1 Northern Rivers Qld. V. 348,
No 105, 1880 F424
Ambassidae 417 Pseudoambassis castelnaui Macleay C 4 Murrumbidgee V. 339, No 79,
1880 F427
Ambassidae 417 Ambassis elevatus Macleay C 6 Endeavour River V. 338, No 75, 1880
F432
Ambassidae 417 Ambassis jacksoniensis Macleay C. Approx. 20 Port Jackson V. 340,
No 81, 1880 F434
ee 417 Pseudoambassis ramsayi Macleay H 1 Port Jackson V. 340, No. 80, 1880
436
Ambassidae 417 Ambassis papuensis All and Macleay C 2 Hall Sound, New Guinea 1875
I. 266, pl. 5, fig. 4, 1875 F438
Latidae 422 Pseudolates cavifrons All and Macleay C 3 Port Darwin I. 262, pl. 3, 1875
F442 (a)
Latidae 422 Lates darwiniensis Macleay H 1 Port Darwin II. 345, No 2, 1877 F442(b)
Oligoridae 424 Oligorus gibbiceps Macleay H 1 Murrumbidgee (Yass) X. 267, 1885 F443
Pp. J. STANBURY 207
Epinephelidae 426 Murrayia jenkinsi Macleay H 1 Murrumbidgee (nr. Yass) X. 268,
1885 F448(a) —
Serrenidae 427 Serranus goldiei Macleay H 1 Port Moresby VII. 226, No 9, 1882 F477
Serranidae 427 Serranus alatus All and Macleay H 1 Hall Sound, New Guinea “Chevert”’
Exp. 1875 I. 264, pl. 4, fig. 2, 1875 F482
Serranidae 427 Serranus guttulatus Macleay H 1 Port Jackson III. 33, 1878 F1166
Serranidae 427 Dules haswellii Macleay C 3 Rockingham Bay V. 359, No 126, 1880 F908
Pseudochromidae 431 Cichlops filamentosus Macleay C Approx. 10 Port Darwin V. 570,
No 428, 1880 F497
Pempheridae 434 Pempheris macrolepis Macleay C. 2 King George’s Sound V. 516,
No 323, 1880 F503
Lutjanidae 441 Genyoroge bidens Macleay C 2 Port Moresby III. 230, 1882 F524
Lutjanidae 441 Mesoprion obscurus Macleay H 1 Endeavour River V. 331, No 64, 1880
F527
‘Lutjanidae 441 Genyoroge unicolor All and Macleay C 2 Percy Is. “Chevert” Exp. 1875
I. 266, pl. 4, fig. 1, 1875 F528
- Lutjanidae 441 Mesoprion roseigaster Macleay H 1 Rockingham Bay V. 331, No 65, 1880
F530
Pomadasidae 444 Diagramma papuense Macleay C 4 Port Moresby VII. 237, No 49, 1882
F535
Pomadasidae 444 Diagramma multivittatum Macleay C 2 Port Darwin II. 349, pl. 7, fig. 2,
1877 F545
Pomadasidae 444 Diagramma crassilabre All Bae Macleay H 1 Hall Sound New Guinea
1875 I. 271, pl. 5, fig. 5, 1875 F546
Theraponidae 445 Therapon truttaceus Macleay H 1 Endeavour River V. 366, No 141,
1880 F573 .
Theraponidae 445 Therapon truttaceus Macleay (?) 1 Endeavour River V. 366, No 141,
1880 F573 (a)
Lethrinidae 447 Lethrinus punctulatus Macleay C 6 Port Darwin II. 351, pl. 8, fig. 2,
1877 F584
Lethrinidae 447 Lethrinus laticaudis All and Macleay H 1 Percy Is. “‘Chevert’” Exp. 1875
Iee2Gx pls Sates 2, 185) E589
Lethrinidae 447 Lethrinus papuensis All and Macleay H 1 Hall Sound, New Guinea
“Chevert” Exp. 1875 I. 276, pl. 8, fig. 1, 1875 F590
Lethrinidae 447 Lethrinus fusciceps Macleay H 1 Port Darwin II. 350, pl. 8, fig. 1, 1877
F596
Lethrinidae 447 Lethrinus aurolineatus Macleay C 2 Port Moresby VII. 247, No 108,
1882 F1159 :
Girellidae 451 Pachymetopon squamosum All and Macleay H 1 Hall Sound, New Guinea
“Chevert” Exp. 1875 I. 275, pl. 9, fig. 1, 1875 F609
Girellidae 451 Girella elevata Macleay H 1 Port Jackson V. 408, No 235 1880 F1155
Gerridae 458 Gerres longicaudus All and Macleay C 4 Cape Grenville “Chevert” Exp. 1875
22a apie, ieee 2s lo as ENG
Gerridae 458 Gerres cheverti All and Macleay C 2 Cape Grenville “Chevert” Exp. 1875
1, WA, Olle 5 tiles aly ALS e/Gy) Teale
Gerridae 458 Gerres profundus Macleay C 2 Port Darwin II. 350, pl. 7, fig. 3, 1877
F618
Gerridae 458 Gerres carinatus All and Macleay C 2 Darnley Island “Chevert” Exp. 1875
I. 273, pl. 7, fig. 4, 1875 F620
Gerridae 458 Gerres bispinosus All and Macleay C 2 Hall Sound, New Guinea “Chevert”
Exp. 1875 I. 273, pl. 7, fig. 3, 1875 F621
Scorpidae 481 Scorpis vinosa All and Macleay H 1 Cape York I. 277, pl. 9, fig. 2, 1875
F674
Chaetodontidae 488 Chaetodon aureofasciatus Macleay C 6 Port. Darwin II. 351, pl. 8,
fig. 3, 1877 F686
Chaetodontidae 488 Chaetodon ocellipinnis Macleay H 1 King George’s Sound III. 33,
1D By Tae aly ALS TENN
Scorpaenidae 493 Tetraroge darnleyensis All and Macleay H 1 Darnley Island I. 278,
DINGS tise le 87/5) BiG:
Scorpaenidae 493 Centropogon echinatus Macleay H 1 Endeavour River V. 436, No 296,
1880 F764
Platycephalidae 506 Platycephalus castelnaui Macleay H 1 King George’s Sound V. 587,
No 456, 1880 F786
. Platycephalidae 506 Platycephalus longispinis Macleay H 1 Trawled outside Sydney
Heads IX. 170, 1884 F1163
208 TYPE SPECIMENS IN THE MACLEAY MUSEUM
Pomacentridae 532 Heptadecanthus longicaudis All and Macleay C 3 Cape Grenville
“Chevert” Exp. 1875 I. 348, pl. 15, fig. 3, 1875 F820
Pomacentridae 532 Dascyllus fasciatus Macleay H 1 Port Darwin II. 361, pl. 10, fig. 2,
i 2
ley ane 532 Pomacentrus obscurus All and Macleay C 4 Torres Straits I. 343,
pl. 15, fig. 2, 1875 F826
Pomacentridae 532 Pomacentrus dolii Macleay C 2 Port Jackson VI. 65, No 672, 1881
epamiiae Ga Chaerops notatus All and Macleay H 1 Cape Grenville 1875 I. 344, pl. 16,
fig. 1, 1875 F862(a)
Labridae 536 Trochocopus rufus Macleay C 2 King George’s Sound III. 35, pl. 5, fig. 3,
Baca Samet pallida Macleay H 1 Endeavour River VI. 100, No 739, 1881 F893
Coridae 537 Coris papuensis Macleay H 1 Port Moresby VIII. 275, No 378, 1883 F898
Coridae 537 Coris cyanea Macleay C 2 Port Moresby VII. 588, No 216, 1882 F899
Coridae 537 Labrichthys melanura Macleay C 3 Port Jackson VI. 89, No 719, 1881 F920
Coridae 537 Labrichthys dorsalis Macleay H 1 Port Jackson VI. 87, No 716, 1881 F920(a)
Coridae 537 Labrichthys maculata Macleay H 1 King George’s Sound VI. 89, No 718, 1881
F920(b)
Coridae 537 Labrichthys nigromarginata Macleay H 1 Port Jackson III. 35, pl. 3, fig. 3,
1878 F922
Coridae 537 Platyglossus immaculatus Macleay H 1 Port Darwin II. 362, pl. 10, fig. 1,
1877 F935
Coridae 537 Cheilolabrus magnilabrus All and Macleay Cotypes of genus and species 4
Darnley Island “Chevert” Exp. 1875 I. 345, pl. 16, fig. 2, 1875 F1154
Coridae 537 Labrichthys labiosa Macleay H 1 Port Jackson VII. 88, No 717, pl. 1, fig. 2,
1881 F1158
Sparisomidae 539 Heteroscarus castelnaui Macleay C 2 Port Jackson III. 36, pl. 5, fig. 2,
1878 F952 :
Scaridae 540 Pseudoscarus flavolineatus All and Macleay C 3 Cape Grenville I. 346,
pl. 16, fig. 3, 1875 F881
Scaridae 540 Pseudoscarus frontalis Macleay H 1 Port Moresby VII. 590, No 228, 1882
F901
Scaridae 540 Pseudoscarus goldiei Macleay C 2 Port Moresby VII. 590, No. 227, 1882
F1165
Scaridae 540 Pseudoscarus labiosus Macleay H 1 Port Moresby VII. 591, No 231, 1883
F938
Scaridae 540 Pseudoscarus moresbyensis Macleay C 2 Port Moresby VII. 591, No 232,
1882 F883
Scaridae 540 Pseudoscarus nudirostris All and Macleay H 1 Cape Grenville I. 346,
DLL feel sirens 80
Scaridae 540 Pseudoscarus zonatus Macleay H 1 Port Moresby VII. 591, No 230, 1882
F890
Odacidae 541 Olistherops brunneus Macleay C 2 Port Jackson III. 36, pl. 5, fig. 1, 1878
F957
Odacidae 541 Odax brunneus Macleay H 1 Port Jackson VI. 109, No 759, 1881 F957(a)
Eleotridae 544 Hleotris compressus Macleay H 1 Port Jackson II. 358, pl. 9, fig. 7, 1877
F998
Eleotridae 544 Hleotris elongata Alleyne and Macleay H 1 Darnley Island I. 334,
pl. 13, fig. 1, 1875 F962
Eleotridae 544 Eleotris taeniura Macleay H 1 Low. Is. Torres Straits V. 624, No 534, 1880
F963
Eleotridae 544 Agonostoma darwiniensis Macleay C Approx 10 Port Darwin II. 360,
ONE ely aakeny toh ILS 7A AMAA)
Gobiidae 545 Hleotris compressus Macleay H1 Port Jackson II. 358 pl. 9, fig. 7 1877 F998
Gobiidae 545 Gobius lateralis Macleay C 3 King George’s Sound V. 602, No. 485, 1880 F983
Gobiidae 545 Gobius semifrenatus Macleay C 5 Port Jackson V. 598, No 478, 1880 F984
Gobiidae 545 Gobius scabriceps Macleay C 2 Endeavour River Qld. V. 603, No 487, 1880
F990(a)
Gobiidae 545 Gobius nigripinnis All and Macleay C 6 Palm Is. Torres Straits I. 332,
pl. 12; fic. 2, 1875 W992
Gobiidae 545 Gobius darnleyensis All and Macleay C 3 Darnley Island I. 331, pl. 12,
fig. 1, 1875 F992(a)
Gobiidae 545 Gobius cristatus Macleay C Approx. 10 Port Jackson V. 610, No 500, 1880
F993
P. J. STANBURY 209
Gobiidae 545 Gobiodon verticalis All and Macleay C 8 Endeavour River, Darnley Island
(in coral) I. 338, pl. 12, fig. 4, 1875 F987(b)
Gobiidae 545 Apocryptes lineatus All and Macleay C 8 Gape Grenville I. 332, pl. 12, fig. 3,
F994
Sanne 545 Apocryptes bivittatus Macleay C 4 Port Darwin II. 357, pl. 9, fig. 5, 1877
F994(c¢
anise Ee Gobius maxillaris Macleay H 1 Port Darwin II. 357, pl. 9, fig. 2, 1877 F999
Gobiidae 545 Gobiosoma guttulatum Macleay C Approx 8 Port Darwin II. 357, pl. 9, fig. 6,
1877 F1001
Gobiidae 545 Gobius flavidus Macleay C Original label only: No specimen Port Jackson
V. 602, No 486, 1880 F1156
Gobiidae 545 Gobius gibbosus C 5 Endeavour River V. 601, No 485, 1880 F1157
Callionymidae 561-2 Callionymus lateralis Macleay H 1 Port Jackson V. 628, No 543,
1880 F1015
Callionymidae 561-2 Callionymus calcaratus Macleay C 6 Port Jackson V. 628, No 542,
1880 F1016
Opistognathidae 579 Opistognathus SEA TaCES Macleay C 3 Port Jackson V. 570,
No 422, 1880 F1019
Opistognathidae 579 Opistognathus darwiniensis Macleay C 3 Port Darwin II. 355,
pl. 9, fig. 3, 1877 F1020
Opistognathidae 579 Opistognathus maculatus All and Macleay H(?) 1 Torres Straits
“Chevert” Exp. 1875 I. 280, pl. 9, fig. 3, 1875 F1021
Clinidae 585 Lepidoblennius marmoratus Macleay C 3 King George’s Sound III. 34,
pl. 3, fig. 2, 1878 F1025
Clinidae 585 Lepidoblennius geminatus Macleay H 1 Port Jackson VI. 13, No 571, 1881
F1026
Clinidae 585 Cristiceps pictus Macleay H 1 Port Jackson VII. 25, No 589, 1881 F1029
Clinidae 585 Oristiceps fasciatus Macleay H 1 Port Jackson VI. 19, No. 579 1881 F1030
Blenniidae 589 Blennius castaneus Macleay H? 1 Port Stephens (should be Port
Jackson) VI. 5, No 550, 1881 F1034
Blenniidae 589 Salarias filamentosus All and Macleay H 1 Cape York I. 337, pl. 14,
fig. 1, 1875 F1035
Blenniidae 589 Petroscirtes guttatus Macleay C 2 Port Jackson VI. 9, No 557, 1881 F1037
Blenniidae 589 Petroscirtes wilsoni Macleay H 1. Port Jackson IX. 170, pl. 1, 1884
F1037 (a)
Blenniidae 589 Petroscirtes cristiceps Macleay C 4 Port Jackson VI. 9, No 559, 1881
F1037 (b)
Blenniidae 589 Petroscirtes fasciolatus Macleay C Approx. 10 Port Jackson VI. 8,
No 556, 1881 F1038
Blenniidae 589 Petroscirtes rotundiceps Macleay C 2 Port Jackson VI. 9, No 558, 1881
F1041
Blenniidae 589 Salarias irroratus All and Macleay H 1 Low Is. Torres Straits I. 337,
pl. 13, fig. 4, 1875 F1042
Blenniidae 589 Salarias spaldingi Macleay C Approx. 10 Port Darwin II. 358, pl. 9,
fig. 4, 1877 F1045
Blenniidae 589 Salarias atratus Macleay C 2 Port Moresby VII. 361, No 170, 1882
F1047
Blenniidae 589 Salarias geminatus All and Macleay C 2 Torres Straits I. 336, pl. 13,
fig. 3, 1875 F1050
Blenniidae 589 Salaris cristiceps All and Macleay H 1 Darnley Island 1875 I. 338,
pl. 14, fig. 3, 1875 F1050(b)
Blenniidae 589 Salarias auridens All and Macleay H 1 Darnely Island, Torres Straits
I. 338, pl. 14, fig. 2, 1875 F1051
Blenniidae 589 Salarias cheverti Macleay C Approx. 10 Darnley Island VI. 12, No 570,
1881 F1051(a)
Blenniidae 589 Salarias lineolatus All and Macleay H(?) 1 Port Darwin I. 336, pl. 13,
fig. 2, 1875 F1052(a)
Monacanthidae 620* Monacanthus fuliginosus Macleay C 2 Port Moresby VII. 596,
No 261, 1882 F1084
Monacanthidae 620* Monacanthus nigricans Macleay H 1 Port Moresby VII. 596, No 260,
1882 F1087
Monacanthidae 620* Monacanthus guttulatus Macleay H 1 King George’s Sound III. 37,
pl. 4, fig. 2, 1878 F1092
* Now called Aleuteridae.
210 TYPE SPECIMENS IN THE MACLEAY MUSEUM
Monacanthidae 620* Monacanthus macrurus Macleay H 1 Port Jackson VI. 330, No 1029,
1881 F1098
Monacanthidae 620* Monacanthus guntheri Macleay C 2 Port Jackson VI. 314, No. 998,
1881 F1161
Tetraodontidae 624 Tetraodon fasciatus Macleay C Approx. 8 Port Darwin II. 365,
pl. 10, fig. 5, 1877 F1138
Antennariidae 629 Antennarius asper Macleay H 1 Darnley Island V; 580, No 442, 1880
F1146
Acknowledgements
Much of this list was compiled by the previous curator of the Macleay
Museum, Mrs. J. E. Anderson, to whom I am deeply indebted. Mrs. Anderson
labelled the types, which were not designated by Macleay, either Cotype or
Holotype and I have used her suggestions in the list. I am grateful to Dr.
F. H. Talbot and Mr. G. P. Whitley who checked the list and made many
helpful suggestions. I am also grateful to Miss K. J. Smith who spent many
hours checking the references. The errors that remain are my responsibility.
References
ANDERSON, J., 1965.—The Macleay Museum at the University of Sydney. Aust. nat. His.,
15: 47-51.
FLETCHER, J. J., 1920.—The society’s heritage from the Macleay’s. Proc. Linn. Soc.
N.S.W., 45: 185-272.
, (and WALKom, A. B.), 1929.—The society’s heritage from the Macleays, Part II.
Proc. Linn. Soc. N.S.W., 54: 185-272.
MaAcMitian, D. N. S., 1957.—‘“‘A Squatter went to Sea.” Currawong Publishing Company,
Sydney.
WHITLEY, G. P., 1957.—List of type-specimens of recent fishes in the Australian museum,
Sydney. Roneo’d, foolscap: i-iii & 1-40.
REPLACEMENT NAME FOR THE PREOCCUPIED GENUS NAME
ODINIA PERRIER, 1885 (ECHINODERMATA: ASTEROIDEA)
ALAN J. DARTNALL,* Davin L. Pawson,j EvizasetH C. Porsz,t and
Brian J. Smirug
[Read 26th June, 1968]
It was recently brought to the attention of one of us (E.C.P.) that the
generic name Odinia Perrier, 1885 (Echinodermata: Asteroidea) is a junior
homonym of Odinia Robineau-Desvoidy, 1830 (Insecta: Diptera).
We propose here a new name for the asteroid:
Subclass Asrrromea de Blainville, 1830
Order ForcipuLatipa Perrier, 1884
Family BrisincGipag Sars, 1876
Novodinia nom nov. pro Odinia (preoccupied)
Type-species: Odiua semicoronata Perrier, 1885, by subsequent designa-
tion of Fisher, 1917.
Etymology: Novodinia is of feminine gender, derived from the original
name.
Remarks: Novodinia is a widely distributed genus, being represented in
deep water in the Atlantic, Pacific and Indian oceans. In the Southern
Hemisphere two species are known, NV. australis (H. L. Clark) from off
Victoria, Australia, and N. novazealandiae (H. E. S. Clark) from off
the Chatham Islands, New Zealand.
Acknowledgement
The authors’ thanks are due to Mr. David McAlpine of The Australian
Museum who drew attention to the existence of the genus Odinia in the
Diptera.
References
Prrrier, E., 1885. —Ann. Sci. nat., ser. 6, vol. 19(8): 9.
ROBINEAU-DEsvolipy, A. J. B., 1830.—Mem. Presentes Acad. Roy. Sci. Inst. France 2: 648.
*A. J. Dartnall, Department of Invertebrates, Tasmanian Museum, Hobart,
Tasmania.
+D. Pawson, Division of Echinoderms, U.S. National Museum, Smithsonian
Institution, Washington, D.C., U.S.A.
t Elizabeth C. Pope, Department of Echinoderms, The Australian Museum, Sydney,
New South Wales.
§B. J. Smith, Invertebrate Department, The National Museum of Victoria,
- Melbourne, Victoria.
PROCEEDINGS OF THH LINNEAN Society or NEw SoutH WALES, Vou. 93, Part 2
PERMIAN FAUNAS AND SEDIMENTS FROM THE SOUTH
MARULAN DISTRICT, NEW SOUTH WALES
R. E. Wass and I. G. GouLp
Department of Geology and Geophysics, University of Sydney
(Plates xiv—xv)
[Read 26th June, 1968]
Synopsis
A richly fossiliferous Permian outlier is described from the South Marulan District.
Fauna and flora consist of 41 species of which one, Hlimata prima is new. Correlation
of these sediments with beds situated about the Wandrawandian Siltstone-Nowra Sand-
stone boundary in the South Coast Permian sequence is suggested.
The spatial arrangement of pelitic sediments containing leaves, and sandstones
containing a marine fauna enables interpretation of the beds as lagoonal and littoral
shoreline deposits which are associated with a Permian transgression. Alternatively,
they may represent a simple stratigraphic sequence.
INTRODUCTION
The earliest workers in the Marulan-South Marulan district, approxi-
mately 120 miles south-west of Sydney, were Woolnough (1909) and Craft
(1931). Studies made by Osborne (1931, 1949) and Osborne and Lovering
(1952) considered petrological aspects of the batholithic and related rocks.
Additional unpublished results are included in theses by Hind (1950),
Lovering (1950) and Svenson (1950). Woolnough (1909, p. 786) and Craft
(1931, Text-fig. 2) record the small areas of Permian sediments overlying
older rocks. The most recent geological map of the area, the Wollongong
1: 250,000 Geological Series Sheet SI56-9 (2nd edn) has been compiled by
the Geological Survey of New South Wales. This shows the south-westerly
extent of Permian strata referred to the Megalong Conglomerate and undif-
ferentiated Berry Formation to be approximately 4 miles north-east of
Permian sediments described herein. One or two isolated outcrops also occur
on the Ordovician-Devonian contact approximately 14 miles north-east of
South Marulan.
Pertinent differences can be observed between this map and the geology
represented on Text-fig. 1, the latter resulting from a study by Gould (1966).
The Silurian acid volcanics and interbedded sediments are separated from
Devonian batholithic rocks. On the Wollongong 1: 250,000 geological map
the distribution of these units suggests that they are not completely separated.
Another major difference is that the fault separating Ordovician and Silurian
strata near South Marulan is interpreted here as an unconformity.
GENERAL PALAEOZOIC SRATIGRAPHY
The lowest stratigraphic unit is the Tallong beds. They are composed
of an undifferentiated sequence of isoclinally folded slates, quartzites and
phyllites considered by Sherrard (1949) to be late Upper Ordovician in age.
Unconformably overlying the Tallong beds is the Bungonia limestone.
The basal portion is massive limestone but this becomes increasingly
arenaceous towards the top. Because of the associated fauna, Favosites
PROCEEDINGS OF THE LINNEAN Society or New SourH WALES, Vou. 93, Part 2
R. E. WASS AND I. G. GOULD 213
gothlandicus and F. allani, Heliolites, Tryplasma, stromatoporoids and
pentamerid brachiopod casts (Svenson, 1950; Flinter, 1950; Gould, 1966) the
unit is regarded as Middle-Upper Silurian in age. Toscanites, tuffs and
tuffaceous labile and sublabile sandstones with minor pelites previously
included in the batholithic rocks are named the Tangerang volcanics. On
field evidence they are considered to overlie conformably the Bungonia lime-
stone and to be Upper Silurian in age, pena extending into ne Lower
Devonian.
The Glenrock granodiorite, a term used by Woolnough (1909) for a
major component of the Marulan Batholith, intrudes the three units already
discussed. Naylor (1939) and Browne (1950) assign to the granodiorite a
probable late Middle Devonian and a Tabberabberan age respectively.
Permian sediments are composed of leaf-rich pelites and fine to medium
grained labile and sublabile calcareous sandstones which contain a rich
marine fauna and plant detritus. Their position on the Bungonia limestone-
Tangerang volcanics contact coincides with a major physiographic break
which extends in a southerly direction towards the Shoalhaven River gorge.
They outcrop approximately one mile south-west of South Marulan at Grid
Ref. 70493024 Wollongong 1: 250,000 geological map. Outcrop, which is
mainly rubble, covers a kidney-shaped area of approximately 40,000 square
yards.
The pelites in the southern portion of the outcrop are composed of 95%
clay size particles and brown organic matter, probably leaves. There are
small amounts of quartz, zircon and mica.
The major portion of the body consists of detritus comprising quartz
(45%), feldspar (25%), rock fragments (25%) with accessory muscovite,
biotite and heavy minerals, including hornblende, tourmaline and opaques.
Matrix constitutes up to 5% of the rock. Sorting is fair with grains ranging
in size from 0:05 mm. to 2°55 mm. in mean diameter. Mineral grains are
angular to subangular while most lithic fragments are subrounded to rounded.
Quartz grains cover the entire grain size range and are dominantly of
two types. Some grains show undulose extinction, mosaic domains and often
with included zircon needles, minute dusty inclusions and rare feldspar laths.
The second type commonly exhibits clear extinction and trails of small
inclusions. Fragment shapes often suggest hexagonal peripheries while pseudo-
inclusions and resorbed rims are apparent.
Feldspar grains generally range in mean diameter from 0-4 mm. to
1-0 mm. although rare grains are only 0-05 mm. across. Potassic feldspar
and plagioclase occur in approximately equal proportions. K-feldspar is often
perthitic and may show sericitisation. Rare myrmekitic growths rim some
grains. Some show crosshatched twinning and may be identified positively
as microcline. Plagioclase is more commonly altered to chloritic products.
Colourless muscovite and altered biotite flakes are scattered throughout.
Heavy minerals are accessory and consist of (i) opaques, (ii) amphibole
lath fragments up to 0 mm. long, mostly chloritised and (iii) rare
tourmaline, pleochroic from blue-green to yellow-green.
Lithie fragments generally show extensive alteration and may be
separated into four groups:
(i) acid volcanics showing embayed quartz, feldspar and biotite in a
once glassy mesostasis,
(ii) granitic fragments composed of a mosaic of quartz ne and
perthitic feldspar,
214 PERMIAN FAUNAS AND SEDIMENTS.
1_—— 5 ,
Permian beds. ji GCOLO aes Corie qu MN
Apt ppsee Geo/ogica/ Boyogaries
2g, Glenrock granooiorite. ——— 7©° USRerred)
St? Tangerang volcanics. ______- Roads
ise) Bungonia limestone. ——— Rai/way:
Taliong beds. 2 4s | km.
Fie. 4 _————— iy
Text-fig. 1. General Geological Map of the South Marulan area (after Gould, 1966).
R. BE. WASS AND I. G. GOULD 215
(iii) trachytic (?) fragments with abundant feldspar laths defining a
flow foliation in a once glassy ? groundmass, and
(iv) rare and extremely fine grained fragments of slates.
Matrix consists of very fine detrital grains with abundant clay minerals.
Patchy iron staining is prevalent. Post depositional mineralogical change
is evident where areas of complete chloritisation occur. The chlorite forms
intergrowths of radiating spherules. These are colourless to pale yellow and
generally show grey interference colours but are rarely isotropic.
Consideration of detritus present indicates three major sources.
1. The Tangerang volcanics as evidenced by the second type of quartz
discussed and also by recognisable acid volcanic fragments,
2. the batholithic rocks themselves due to the first type of quartz and
granitic fragments, and
3. the regionally metamorphosed Tallong beds as evidenced by slate
fragments and the pleochroic tourmaline, characteristic of these strata.
FAUNA AND FLORA
Fauna and flora identified are listed below.
Leaf detritus is found dominantly in the pelites but some is
sandstone.
The fauna occurs in the
enclosed in the sandstone. In the list, an asterisk (*) indicates the species
is discussed later and a stroke (/) indicates the species is figured.
Cladochonus sp.
Conularia inornata Dana
Conularia cf. tuberculata Fletcher
Fenestella canthariformis
(Crockford)
Fenestella dispersa (Crockford)
Fenestella granulifera
(Crockford )
Polypora woodsi (Etheridge, Jr)
Protoretepora ampla (Lonsdale)
*/Stenopora gracilis (Dana)
Ambikella cf. ingelarensis
(Campbell)
/Ambikella cf. isbelli (Campbell)
Ambikella cf. mantuanensis
(Campbell)
/Ambikella cf. undulosa
(Campbell)
*/Fletcherithyris cf. amygdala
(Dana)
/Fletcherithyris parkesi Campbell
/Gilledia ulladullensis Campbell
*/Notospirifer cf. minutus Campbell
/Strophalosia clarkei Etheridge Sr
Strophalosia clarkei var. minima
Maxwell
Terrakea solida (Etheridge and
Dun)
*/Terrakea sp.
Trigonotreta stokesi Koenig
/Pleurikodonta cf. elegans
Runnegar
Atomodesma (Aphanaia) sp.
*/Conocardium sp.
Aviculopecten subquinquelineatus
McCoy
*/Elimata prima sp. nov.
*/Stutchburia costata (Morris)
Schizodus sp.
Vacunella cf. curvata (Morris)
Merismopteria sp.
Myonia corrugata? Fletcher
Keeneia minor (Fletcher )
Keeneia ocula (Sowerby)
Peruvispira cf. elegans (Fletcher)
Peruvispira trifilata (Dana)
Strotostoma inflata Fletcher
Tribrachiocrinus sp.
Phialocrinus cf. konincki (Clarke)
Glossopteris ampla Dana
Glossopteris sp.
The occurrence of these sediments on the edge of the Sydney Basin
together with the spatial arrangement of leaf-rich pelites and sandstones
_ containing a marine fauna indicated that a shoreline may have extended
over the area. Because of this the surrounding area was surveyed accurately
216 PERMIAN FAUNAS AND SEDIMENTS
to obtain the outcrop pattern and to plot the boundary between the pelites
and sandstones. Using a scale of 1 inch equals 60 feet, a theodolite stadia
traverse established control with sufficient accuracy for a 2 feet contour
interval. This has been increased on the final plan to avoid congestion.
Detail and geological boundaries were obtained using a telescopic alidade and
plane table. The accuracy of the Permian-Silurian boundary to the east may
be doubted due to the movement of Permian talus downslope and the super-
ficial similarity of the Permian sandstone and Bungonia limestone lithologies.
The heavily wooded nature of the terrain also hindered surveying.
The map resulting from the survey appears as Text-fig. 2.
/ Permian sandstones
/ Permian pelites
Tangerang volcanics
Bungonia limestone
4? Traverse Station
Spot Height
AY Topographic Contour
>r— Road
——— Fence
——— Power Line
Geological Boundaries :—
—— Accurate
METRES
FEET
6o ° 60 Teo
LEVEL DATUM ASSUMED
——— Inferred
SS
i FIG. 2 oon Shoreline
Text-fig. 2. Geological and Survey map of the Permian outlier and surrounding
area.
R. E. WASS AND I. G. GOULD 2h
~ DEPOSITIONAL INTERPRETATION
A division between pelites and sandstones is developed in the southern
part of the Permian outcrop along an assumed topographic level of 973 feet
to 979 feet. This is approximately 110 feet below the ridge top of Tangerang
volcanics to the west. The lowest occurrence of Permian sediment is some
130 feet below this ridge. The ridge of Tangerang volcanics to the north and
the eastern ridge formed by the Bungonia limestone are not as high
topographically but enclose the Permian sediments on three sides with a
topographical opening to the south.
In other areas on the edge of the Sydney Basin, the Permian is thin
and in places is abnormal in character. The base of the Permian traces an
irregular junction with older rocks so sediments such as these could have
been deposited on a surface of some relief. It is possible that this area,
affected by a period of Permian transgression contained many inlets and bays
which received sediment transported from an eroding high composed of older
rocks to the west. The South Marulan area may be interpreted ag an inlet
with the sandstones being ina littoral environment and the pelitic sediments
representing a lagoonal or deltaic environment on the edge. The division
between pelites and sandstones could be interpreted then as a shoreline.
Subsequent erosion related to a period of regression in the Permian and
post-Permian erosion would remove some of the sediments giving rise to
the disposition of beds as in Text-fig. 2.
A possible reconstruction of the area in Permian time appears as
Text-fig. 3.
Text-fig. 3. Block diagram showing possible topographic elements during deposition
of South Marulan strata during Permian time.
We do not feel that the deposits can be representative of both
transgression and regression. The marine sediments can be referred to a
transgression but it is unlikely that the pelites are part of a regressive phase
as one would expect sediments associated with such a regression to become
coarser. A regression followed by a transgression is out of the question as
the latter must be invoked to allow marine environment to cover the area.
The possibility that the sandstones may overlie pelites must be recognized.
To the north and east, shorelines are formed and coarse sediments are overlain
by finer sediments with a northerly or north-easterly dip (Craft, 1931). With
such a situation here, the dip would be south-easterly but this would depend
to some extent on associated Permian topography. Unfortunately no dips
can be recorded from the South Marulan sediments.
218 PERMIAN FAUNAS AND SEDIMENTS.
If the marine deposits are littoral in origin, one would expect to find
some evidence of turbulence near the junction with the pelitic sediments and
evidence of quieter conditions some distance from this region. The fauna
close to the supposed shoreline consists entirely of broken polyzoan detritus,
ramose colonies of stenoporids and fenestellids with a large proportion of
leaves. This supports a littoral origin. With increasing distance from the
pelitic sediments, brachiopod and mollusc individuals increase in number
with their valves united. Polyzoan colonies become more complete. Only
near the edge of the littoral deposits is there evidence supporting turbulence.
Elsewhere the fauna seems to have lived in relatively quiet conditions in
shallow water. This would be in keeping with the possible physiographic
position and the palaeogeography.
In considering the palaeogeography of the south-western portion of the
Sydney Basin, one must take into account the study made by Gostin (1968).
His thesis has shown that along the far south-western margin of the Basin
in Permian time, sediments unconformably overlying pre-Permian basement
become younger in a general north-westerly direction. This is based on faunal
and field evidence. In the far south of the Basin, the lowest unit of the
Conjola Formation unconformably overlies basement whereas to the west of
Ulladulla, the topmost unit of the Conjola Formation, and further to the
north-west, the Wandrawandian Siltstone and Nowra Sandstone overlie
basement. In other words, there is a progressive transgressive phase in a
north-westerly direction along the south and south-western margins of the
Basin. The precise direction of the transgression cannot be determined at
this stage due to the lack of satisfactory control points in the Nowra-Berry
district.
Faunal evidence to support this transgression is adequate. Faunas in
the lowest unit of the Conjola Formation are considered by Runnegar (in
press) to be Dalwood equivalent, possibly correlative with the Allandale
Formation. Further to the northwest, younger faunas in the topmost unit
of the Conjola Formation and the Ulladulla Mudstone reveal similarities to
faunas in the lower part of the Branxton Formation in the Hunter Valley.
Therefore, it is not surprising that an analysis of the South Marulan fauna
reveals that it is equivalent to fauna in high Wandrawandian Siltstone-low
Nowra Sandstone. It is probably correlative with high Branxton or low
Muree Formation. This is discussed in more detail subsequently.
Systematic DESCRIPTIONS
Phylum Mo.uuusca
Class PrLEcypopa
Superfamily ?
Family ConocarpiipAE Neumayr
Genus Conocardium Bronn, 1835
Type Species: (by monotypy) Cardium (Conocardium) elongatum
Sowerby, 1812, p. 188, pl. 82, fig. 3.
Diagnosis: See La Rocque, 1950, p. 317.
CONOCARDIUM Sp.
(Pl. xiv, Figs 10-14)
In the description the shell is oriented in the sense of Branson (1942)
and La Rocque (1950) and is the opposite of Fletcher (1943).
Description: Shells are equivalve and small. Anteriorly they are alate
with a flattened area near the hinge line. A characteristic key-hole shaped
ventral gape is developed along more than half the anterior ventral margin
R. E. WASS AND I. G. GOULD 219
which is slightly curved. The region adjacent to the gape on an internal
mould is crenulate with more than 13 crenulations present. They decrease
in size and are more closely spaced posteriorly, and can be traced from the
margin to the umbonal region. Concentric ornament is not developed as
strongly. Carinae are not well developed. The semi-crecentic posterior area
is ornamented with 15 or more radial, and concentric plicae, producing small
inflections on the slightly curved margin. The posterior tube is produced
closer to the umbones than to the posterior extremity. Umbones are small
and centrally situated. The hinge line is straight and long. The anterior
ligament area is long and narrow and the posterior area is large and wide.
On an external mould, 24 primary radial ribs can be traced from the
umbonal region to the anterior ventral margin. This number increases near
the margin as secondary ribs arise between the primaries.
Discussion: This species has morphological resemblances to Conocardium
robustum Fletcher, 1943, because of its strongly inflated carinal area, the
shape of the anterior gape and the oblique nature of the carina. It is much
smaller than this species but this may be due to immaturity. The species
has a more oblique carina than Conocardium australe (McCoy).
Specimens catalogued from South Marulan are S.U.P. (Sydney University
Palaeontological Collection) 12622 A, B, C.
Dimensions: These are related to the long, straight anterior region.
12622 (right valve) lLength...... 27 mm.
Thickness... 10
Superfamily Carpiracna ?
Family MyocHoncHIpAE Newell
Genus Stutchburia Etheridge, Jr, 1900
Type Species: (by original designation) Orthonota ? costata Morris,
1845, p. 273, pl. 11, fig. 1, from the Permian of the Illawarra region, New
South Wales.
Diagnosis: See Dickins, 1963, p. 95.
STUTCHBURIA cosTATA (Morris), 1845
(Pl. xiv, Figs 1-5)
Diagnosis: Shell slightly expanded towards rear; radial plications
confined to posterior portion of the shell.
Description: Shells become slightly higher and elongate posteriorly ;
umbones are not prominent; a long deep ligament groove is placed posterior
to the umbones. There is a great variation in size. Morphology developed is
constant except for the muscle scars. On all specimens the anterior adductor
scar has a prominent posterior buttress. This is higher dorsally and wider
anteriorly. However, on small specimens the anterior region of this scar
overhangs the margin. This is not so with larger specimens in which the
anterior adductor scar is divided into a dorsal one-third and a ventral two-
thirds by a low ridge which is higher dorsally. Posterior scars are nearly
semi-circular with the diameter paralleling the hingeline but they are slightly
asymmetrical towards the anterior. Anterior scars are oval. Elongate, narrow
pedal scars are evident in front of the umbo on all specimens. The pallial
line is entire meeting the anterior scar at its postero-lateral margin and the
_ posterior scar in its ventral region. The hinge is edentulous with a slight
twist to the right in front of the umbones. Ornamentation consists of con-
220 PERMIAN FAUNAS AND SEDIMENTS —
centric growth lines and upwards of 5 strong radial plicae in the posterior
region.
Discussion: The occurrence of a form possessing coarse costae with a
subdivision of the anterior adductor scar by a ridge is interesting as Etheridge
(1900) stated that the coarse costae were characteristic of S. costata whereas
the latter feature was characteristic of S. compressa. Further studies may
show the two species to be identical.
Specimens catalogued are S.U.P. 12610-12614, 12615, A, B, C, 238573
AS IBS C:
Dimensions: (right valve)
Length Height Thickness
23573A 63-0 25.0 9-5 mm.
12610 9-6 6-0 2-5 mm.
12611 5-1 2°9 1-0 mm.
Superfamily PrcTiINAcEA
Family Limipar d’Orbigny
Genus Elimata Dickins, 1963
Type Species: (by original designation) Elimata guppyi Dickins, 1963,
p. 93, pl. 15, figs 6-18, from the base of the Permian, Poole Sandstone,
Western Australia.
Diagnosis: See Dickins, 1963, p. 93.
ELIMATA PRIMA, Sp. NOV.
(Pl. xiv, Figs 6-9)
Holotype: 12607 S.U.P. from the northern section of Permian sediments
at 70493024 Wollongong 1: 250,000 Geological Series Sheet SI 56-9, approxi-
mately one mile south-west of South Marulan, New South Wales.
Diagnosis: Robust, convex, markedly opisthocline shells with a short,
straight hinge line. :
Description: Shells are opisthocline with a short, straight hinge line.
Weakly developed umbonal ridges separate small flattened areas from the
rest of the shell. The posterior ridge is sharper and more distinct than the
rounded anterior ridge which produces a steep slope on the dorsal antero-
lateral region of the shell. Concentric growth lines are present and sometimes
fine radiating plicae can be observed near the ventral margin.
Discussion: This species is larger, more opisthocline and possesses a
shorter hinge line than the type species. This is the first published record
of the genus from the Eastern Australian Permian strata. Dickins (1964)
referred to Hlimata sp. nov. from the Ingelara Formation in the south-
western portion of the Bowen Basin, Queensland.
Specimens catalogued from South Marulan are S.U.P. 12606-12609.
Dimensions: (right valve)
Length Height Thickness
12607 13 12 2-0 mm.
12609 13+ 15 2-5 mm.
Phylum BracHIOPpoDpA
Class ARTICULATA
Order TEREBRATULIDA
Family Dirtasmatipa® Schuchert
Genus Fletcherithyris Campbell, 1965
R. E. WASS AND I. G. GOULD
i)
bo
hk
FLETCHERITHYRIS cf. AMYGDALA (Dana), 1847
(Pls Seep Tey P(e Lb)
Remarks: Specimens assigned to this species show some variation from
those described by Campbell. One has a characteristic V-shaped septalium
with growth lines normal or directed slightly posteriorly to its length. This
is different from FF’. amygdala’ (Dana) where they are directed mainly
anteriorly (Campbell, 1965, pl. 6, figs 24, 32) and F. farleyensis Campbell
in which they are directed posteriorly (Campbell, 1965, pl. 6, ‘fig. 4).
The septalium developed is of similar length to amygdala and similar to
farleyensis in height. The lateral commissure is intermediate between these
two species (Campbell, 1965, pl. 6, figs 7, 29).
A specimen smaller than the normal. farleyensis may be I’. farleyensis
faba Campbell but the characteristics anterior flattening of the pedicle valve
in the latter species is not conspicuous. Growth lines on the septalium are
~ normal to its length.
Specimens catalogued are S8.U.P. 25557-25558.
Dimensions:
Length Width Height
25557 31 22 13-5 mm.
25558 21 14 8-0 mm.
Order SPirRIFERIDA
Family Martinipar Waagen
Genus Notospirifer Harrington, 1955
Type Species: (by original designation) Spirifer darwini Morris, 1845,
p. 279 from the Permian, ? Muree Formation at Glendon, Hunter Valley,
New South Wales.
Diagnosis: See Campbell, 1959, p. 342; Waterhouse and Vella, 1965, p. 70.
Norosprrirer cf. MINuTUS Campbell, 1960
(Pl. xv, Figs 1-5)
Remarks: Specimens from South Marulan show much variation in
external morphology. In none could the fold in the brachial valve be
described as being flat on top. There is a gradation from folds with a
shallow sinus to folds with a strong sinus developed. Specimens with the
latter feature are considered to be gerontic, this being based also on the
nature of the plications. The sulcus of the pedicle valve can have a small
fold developed along its midline. In mature specimens, the number of
plications is the same in N. minutus but they are more strongly developed.
In younger specimens the greatest width is more posterior than in older
specimens.
Internally the ventral adminicula are similar to those figured by Campbell
(1960, pl. 140, fig. 7) and only in small specimens do they tend to become
subparallel posteriorly. Dorsal adminicula are shorter than the ventral and
widely divergent. They are noticeably longer on specimens with five well
developed plicae. .
Specimens catalogued from South Marulan are S.U.P. 25544, 25549-52,
25555, 25560, 25561, 25565, 25572, 25573, 25576.
Dimensions: (brachial valve)
Length Width Height
25551 15 28 6-8 mm.
25552 13 23 9:0 mm.
25572 20 36 8-0 mm.
25573 20 39 10:0 mm.
222, PERMIAN FAUNAS AND SEDIMENTS
Order STROPHOMENIDA
Family Linopropuctipag Stehli
Genus Terrakea Booker, 1930
Type Species: Productus brachythaerus Morris, 1845, from the Permian
of New South Wales. (See Maxwell, 1956, Heming, 1957.)
Diagnosis: See Moore, 1965, p. H503.
TERRAKEA Sp.
(Re sxave ie Seg)
Remarks: Fragmentary remains from South Marulan reveal a wide,
straight hinge line. Umbonal shoulders are not steep and the umbo which is
blunt and not strongly thickened only slightly overhangs the hinge line.
Diductors are longitudinally striated; adductors are finely dendritic.
Features of the umbonal region indicate that this species is remarkably
similar to Terrakea sp. from the Ingelara Formation and Catherine Sandstone
of the Springsure 1: 250,000 Sheet area of Queensland. The blunt nature of
the umbo with an absence of strong thickening enables a separation from
Terrakea solida which is found generally at a higher stratigraphic horizon.
Specimens catalogued from South Marulan are 8.U.P. 25540, 25545.
Phylum Poryzoa
Class GYMNOLAEMATA
Order TREPOSTOMATA
Family StreNoporipAE Waagen and Wentzel
Genus Stenopora Lonsdale, 1844
Type Species: (by subsequent designation of Ulrich, 1890, p. 375)
Stenopora tasmaniensis Lonsdale, 1844, p. 178 from the Permian of Southern
Tasmania.
Diagnosis: Zoarium massive, ramose, encrusting, bilaminar; zooecial
tubes thin walled in axial region and exhibiting a definite annulated habit
in the peripheral region; diaphragms absent; apertures oval or rounded,
mesopores and acanthopores present; maculae and monticules may be
developed.
STENOPORA GRACILIS (Dana), 1849
(Text-fig. 4)
Diagnosis: Ramose zoarium with a narrow peripheral region and two or
three rows of annulations; mesopores not well developed, acanthopores
numerous; maculae irregular.
Text-fig. 4. Oblique tangential section through Stenopora gracilis, 16431, x 10.
Remarks: Specimens from South Marulan possess ramose zoaria with a
diameter of 1-0-5-0 mm. The peripheral region is 1:2 mm. wide in a zoarium
of diameter 5-0 mm. Zooecial tubes leave the axis at an angle of 35°-40° and
in the peripheral region they are normal to the periphery.
R. E. WASS AND I. G. GOULD 223
The small, ramose zoaria and the nature of the peripheral region are
considered adequate to include these specimens in Stenopora gracilis. The
species, S. nigris Crockford is similar to S. gracilis. According to Crockford
(1943) both are charaterised by a narrow, peripheral region with two rows
of annulations, numerous acanthopores and few mesopores. S. gracilis is
the finer species but an examination of S.U.P. material reveals that measure-
ments are more similar to S. nigris than those stated by Crockford. |
Specimens catalogued are 8.U.P. 16431-16438.
AGE AND CORRELATION
; Recent work by Campbell (1965), Dickins, Gostin and Runnegar (in
press), Gostin (1968) and Runnegar (in press) together with studies made
during this decade in the Permian of Queensland, particularly by Dickins
(in press) enable a reasonable correlation and stratigraphic analysis of the
South Marulan fauna to be made with similar faunas on the South Coast
and Hunter Valley in New South Wales and with the Bowen Basin in
Queensland. Many species are long ranging and of little value. A conspicuous
feature of the fauna is the absence of genera such as Taeniothaerus,
Grantonia, Deltopecten and Hurydesma and particular species of the genera
Notospirifer, Cancrinella, Terrakea and Ambikella. This immediately indi-
cates that the fauna is younger than Fauna II in the Bowen Basin and
similar faunas elsewhere. However, species such as Keeneia ocula, Gilledia
ef. ulladullensis, Fletcherithyris parkesi, Ambikella cf. isbelli and A. cf.
undulosa are recorded from the Conjola Formation, partly equivalent to
Fauna II (Dickins e¢ al., in press).
The use of species of Ambikella for correlative purposes in the South
Coast and Hunter Valley Permian sequences has limited value as most
species were originally described from Queensland and those in the Sydney
Basin are in urgent need of critical examination. Therefore, all species of
Ambikella from South Marulan are prefixed by cf. It is worthy of note that
the species possess a shallow sulcus and the adminicula are elongated in the
brachial valve. These are considered to be features of Ambikella spp. in
Fauna III of the Bowen Basin.
Species from South Marulan regarded as significant and of use in
correlation are Fletcherithyris cf. amygdala, Notospirifer cf. minutus,
Terrakea sp., T. solida, Plewrikodonta cf. elegans and Vacunella cf. curvata.
The occurrence of EHlimata prima, sp. nov. may be of some value as is the
absence of the terebratuloid Marinurnula. In the southern region of the
Sydney Basin the absence of this genus is of stratigraphic significance as
it indicates that the fauna is not younger than the Nowra Sandstone
(Campbell, 1965).
Of the South Marulan species, Dickins et al. (in press) record
Strophatlosia cf. clarkei, Terrakea sp. and Fletcherithyris amygdala from the
Wandrawandian Siltstone and Vacunella cf. curvata, Notospirifer cf. minutus,
Strophalosia cf. clarkei and Ambikella cf. isbelli from Nowra Sandstone.
Of the other species considered significant, Pleurikodonta elegans has been
recorded by Runnegar (1965) from Fauna IV and possibly high Fauna ITI
in the Bowen Basin. This is the first record of the genus outside this area.
Atomodesma (Aphanaia) sp. has been identified from the Oxtrack Formation
(low Fauna IV) and in the Springsure area, Hlimata occurs first in the
Ingelara Formation (middle Fauna III) and in strata containing a high
- Fauna III there is an incoming of Terrakea sp. and Notospirifer cf. minutus.
The boundary between the Nowra Sandstone and the Wandrawandian Silt-
224 PERMIAN FAUNAS AND SEDIMENTS
stone may be equivalent to the upper part of the Gebbie Subgroup (high
Fauna III) in the Bowen Basin. It seems therefore, that the South Marulan
strata may be best correlated with the strata about the Wandrawandian
Silstone-Nowra Sandstone boundary. Considering the Hunter Valley sequence
in terms of the South Marulan fauna, correlation with the upper part of the
Branxton Formation or low Muree Formation is suggested.
Terrakea solida occurs only in Fauna IV. Aviculopecten subquinque-
lineatus, Keeneia minor, Stutchburia costata, Gilledia ulladullensis, Fletcheri-
thyris parkesi and Fenestella canthariformis together with species of
Ambikella appear high or low in the sequence and it seems that correlatives
of Fauna III in the southern Sydney Basin contain species found in Faunas
II and IV as well as Fauna III of the Bowen Basin. Dickins e¢ al. (in press)
have already recognised this feature and consider that it may be due to
either an hiatus during Fauna III time in the South Coast sequence or to
the faunas appearing at slightly different times due to geographical and
environmental factors.
A complete faunal study of the South Coast Permian sequence, especially
the productids, spiriferids and pelecypods would be beneficial and as a result,
correlation of the South Marulan strata with the South Coast sequence may
need revision.
Acknowled gments
For their interest in this study and for critical discussion on pertinent
portions of the manuscript we wish to thank Dr. J. M. Dickins, Bureau of
Mineral Resources, Canberra, Dr. K. S. W. Campbell and Mr. V. A. Gostin,
Department of Geology, Australian National University, Canberra and Dr.
B. N. Runnegar, Department of Geology, University of New England,
Armidale. Dr. B. D. Webby has advised us regarding interpretation of the
deposit and editorial matters. Technical assistance has been rendered by Mr.
B. R. Lambert, Mr. F. Smith and Mr. G. Z. Foldvary. The study has been
financed by a University of Sydney Research Grant. One of us (I.G.G.)
acknowledges assistance from the New South Wales Department of Mines.
References
Booker, F. W., 1930.—A review of some of the Permo-Carboniferous Productidae of
New South Wales with a tentative reclassification. J. Proc. r. Soc. N.S.W., 64: 65-77.
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Browne, W. R., ed., 1950.—The Geology of the Commonwealth of Australia. 3 vols,
Edward Arnold and Co., London. :
CAMPBELL, K. S. W., 1959.—The Martiniopsis-like Spiriferids of the Queensland Permian.
Palaeontology, 1(4): 333-350.
, 1960.—The brachiopod genera Ingelarella and Notospirifer in the Permian of
Queensland. J. Paleont., 34(6): 1106-1123.
, 1965.—Australian Permian Terebratuloids. Bull. Bur. miner. Resour. Geol.
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Crart, F. A., 1931.—The physiography of the Shoalhaven River Valley. 1. Tallong-
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CrockForp, J., 1943.—Permian Bryozoa from eastern Australia, Part III; Batostomellidae
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Dawa, J. D., 1847.—Descriptions of fossils from Australia. Am. J. Sci., 54: 151-160.
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Dicxins, J. M., 1963—Permian Pelecypods and Gastropods from Western -Australia.
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miner. Resour. Geol. Geophys. Aust., 1964/27 (unpubl.).
R. HB. WASS AND I. G. GOULD 225
, (in press);—Correlation of the Permian of the Hunter Valley, New South
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Aust.: 80.
, Gostin, V. A., and Runnecar, B. (in press).—Correlation and age of the
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Frintrer, B., 1950.—The geology of the Bungonia area. Unpubl. Hons Thesis, Univer-
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Gostin, V. A., 1968.—Lower Permian Sediments, southern extremity, Sydney Basin,
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Hemine, F., ed., 1957—Opinion 486. Int. Comm. zool. Nom., 17(8): 105-118.
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EXPLANATION OF PLATES
PLATE XIV
Stutchburia costata
Fig. 1. 23573A, internal mould of a right valve, x1. Fig. 2. 23573C, external cast
of ornament, x1. Fig. 3. 23573A, internal mould showing hinge line, x1. Fig. 4. 12611,
internal mould of right valve showing overhang of anterior adductor scar, x 8. Fig. 5.
23573A, internal mould of left valve showing ridge in anterior adductor scar, pedal
retractor and pallial line, x 3.
Cc
226 PERMIAN FAUNAS AND SEDIMENTS
Hlimata prima, sp. Nov.
Fig. 6. 12607, internal mould of left valve, x2. Fig. 7. 12606, internal mould of
right valve, x2. Fig. 8. 12609, internal mould of right valve, x2. Fig. 9. 12607, side
view of internal mould of left valve, x 2.
Conocardium sp.
Fig. 10. 12622C, external cast of ornament on right valve showing secondary ribs
developing between primaries, x2. Fig. 11.12622A, internal mould of right valve, x 1-5.
Fig. 12. 12622A, internal mould showing keyhole shaped ventral gape, x1-5. Fig. 13.
12622A, umbonal view of internal mould showing straight hinge line and posterior tube,
x15. Fig. 14. 12622A, posterior view of internal mould showing position of posterior
tube and inflections on margin produced by plicae, x 1:5.
Pleurikodonta cf. elegans
Fig. 15. 12624A, anterior view of internal mould, x2. Fig. 16, 12624B, external
mould showing crenulate commissure, x3. Fig. 17. 12616, internal mould showing
megadesmatid tooth fold, x4.
Strophalosia cf. clarkei
Fig. 18. 25539, internal mould of pedicle valve, x 1:5.
Terrakea sp.
Fig. 19. 25540, oblique view of umbonal region on internal mould of pedicle valve, x 1.
PLATE Xv
Notospirifer cf. minutus
Fig. 1. 25572, internal mould of brachial valve, x2. Fig. 2. 25551, posterior view
of internal mould, x2. Fig. 3. 25572, posterior view of internal mould showing sinus
in fold of brachial valve, x2. Fig. 4. 25552, internal mould of pedicle valve, x2. Fig. 5.
25551, internal mould of anterior commissure, x 2.
Fletcherithyris parkesi
Fig. 6. 23452, internal mould of brachial valve, x1-:5. Fig. 12. 23452, internal mould
of anterior commissure, x 1-5.
Fletcherithyris cf. amygdala
Fig. 7. 25557, lateral commissure on internal mould, x1:5. Fig. 11. 25557, internal
mould of brachial valve, x 1:5.
Gilledia ulladullensis
Fig. 8. 25559, internal mould of brachial valve, x1-5. Fig. 9. 25559, internal mould
of pedicle valve, x1-5. Fig. 10. 25559, anterior commissure on internal mould, x 1:5.
Fig. 13. 25559, lateral commissure on internal mould, x 1-5.
Ambikella cf. isbelli
Fig. 14. 23451, anterior commissure on internal mould, x1. Fig. 15. 23451, internal
mould of pedicle valve, x1:5. Fig. 16. 23451, posterior view of internal mould, x1°5.
Ambikella cf. undulosa
Fig. 17. 25579, internal mould of pedicle valve, x1.
Proc. Linn. Soc. N.S.W., Vol. 93, Part 2 PLATE XIV
xu
eo
Hoy eae
Sanit
ON THE FIRST OCCURRENCE OF A CLIMACOGRAPTUS BICORNIS
WITH A MODIFIED BASAL ASSEMBLAGE, IN AUSTRALIA
Henry Moors
University of Melbourne
[Read 31st July, 1968]
; Synopsis
This paper describes the first Climacograptus bicornis with basal spines modified
by the presence of accessory spines, found in Australia, at Tallong, N.S.W. It is com-
pared with a more complicated species Cl. venustus found in China and its position
with respect to the evolution of Cl. venustus from Cl. bicornis discussed. The structure
of the basal assemblage is described more fully than, and compared with, the Chinese
species and its possible function discussed.
INTRODUCTION
In 1959, S. C. Hsu (Hsu, 1959) described a new species of Climacograptus
with a peculiar basal modification from northwestern Hupeh, China, naming
it Cl. venustus and showed a possible line of evolution of the species from
and through other recognized species. Hsu states (op. cit., p. 351) “It appears
hardly possible that the complicate appendage consisting of the powerful
principal spines with fully developed accessory ones could have developed
directly from the simplest form with only two thin lateral spines. There must
be some forms intermediate between them, which have not yet been discovered
at present.” He then showed how Cl. venustus could have been developed
from Cl. bicornis and backed this line of descent by an analogy with the
line of descent of Cl. peltifer from Cl. bicornis. However, Hsu had only two
specimens, both of the same degree of development, and so could only make
a guess at the evolution of his species. The finding of this specimen at Tallong,
N.S.W., may help to clarify the evolutionary trend as it appears to be of
intermediate development between Cl. venustus and Cl. bicornis.
SYSTEMATICS
Family DirpLocraptipaAkn Lapworth
Genus Climacograptus Hall
Climacograptus bicornis Hall subspecies.
DESCRIPTION
The polypary has a minimum length of 20-7 mm., excluding spines, but
would have reached a greater length as the specimen studied was incomplete.
The width at the sicula is only 0-8 mm. and the rhabdosome expands con-
stantly to a maximum of 1-8 mm. at about 1 cm., after which it decreases
gradually to 1-6 mm. distally. A virgula, or median septum, can be seen to
run through the middle of the rhabdosome after the fourth thecal pair.
The thecae are typically climacograptid and appear to be slightly intro-
torted causing the aperture to be introverted into an apparently very shallow,
small excavation, about 1/6th the width of the rhabdosome. Proximally the
thecae number about 13 in 10 mm. with a length of about 0-7 mm., a width
PROCEEDINGS OF THH LINNEAN Sociery or NEw SoutH WALES, VoL. 93, Part 2
228 CLIMACOGRAPTUS BICORNIS WITH A MODIFIED BASAL ASSEMBLAGE
of 0-3 mm. and overlap each other by about 1/3 of their lengths, while distally
they number just under 10 in 10 mm. and are 1:7 mm. in length.
No sicula could be seen, but the proximal end is typified by two large
horizontal spines with a single accessory spine on each. The principal spines
(in the terminology of Hsu, 1959) are both crescentric and together form a
parabolic curve. One is over 43 mm. in length while the other, being
incomplete, measures only 34 mm., and both have maximum widths of approxi-
mately 0:3 mm. About one quarter of the way from the sicula each spine
carries an accessory which is parallel to the rhabdosome, and measures
approximately 1 mm. in length (1:1 mm. and 0-8 mm.); it is of constantly
tapering shape and maximum width of 0-2 mm. The principal spines are
made up of two portions (Fig. 1, a), the main portion being a solid (?)
rod extending for the whole length of the spine and supporting underneath
it a thin tube for about two thirds of its length. The solid rod does not appear
to be affected by the accessory spines.
a b
)
d
[
Fig. 1. Camera lucida drawings of actual specimens from locality 9, Tallong.
(a) The Climacograptus described in this paper, x5. (b) to (f) Cl. bicornis from the
same locality as (a) showing the large range of size of rhabdosome and modifications
of basal spines; all x 23.
This assemblage is markedly asymmetrical, the spines are of different
lengths (one is incomplete but by comparison with the other can be estimated
to be shorter); in this obverse aspect the right hand spine is larger than
the left and also carries an accessory spine larger than that carried by the
other.
DISCUSSION
The specimen here described varies little from both Cl. bicornis and Cl.
venustus appearing to be a natural link between the two.
The nature of the principal spines is apparently clearer here than in
Hsu’s two specimens, and the relationship of the various components better
HENRY MOORS 229
shown. The spines can be seen to consist of a tube of almost constant
diameter supported by a thin rod which continues past its end (see Fig. 1, a)
rather than Hsu’s constantly tapering tube with central rod. Also there
does not appear to be any constriction of the tube in the vicinity of the
auxilliary spines as described by Hsu (an examination of his figures does
show some irregularities in width but not as rhythmic as suggested by his
description, but they could have been badly drawn).
_ The accessory spines appear to differ in some respects from those of Hsu.
Firstly those of Cl. venustus are almost radially disposed on the principal
spines (though less so away from the rhadbosome), Hsu comparing it to a
high toothed cog, while in this specimen the accessory spines lie parallel to
the rhabdosome. Although they are somewhat shorter than in Hsu’s specimens,
this can be explained by their more tangential position on the principal spines
compared to the more radially situated spines of Cl. venustus, as explained
above.
The function of the accessory spines must remain theoretical (see Hsu,
op. cit., p. 351, for some ideas and references) but the suggestion that they
could be modified apertures to allow egress of internal tissue does not seem
valid as the accessory spines end in a very fine point, limiting any aperture
that could be there to very small dimensions, and the principal spines appear
to have apertures of their own, where the supporting rod emerges from its
tube. For the development of the spines of Cl. bicornis see Bulman, 1947, pp.
59-62.
Climacograptus bicornis is a very widespread species, specimens having
been found from probably all the graptolite bearing strata of appropriate
age. It is very diagnostic in appearance and is easily identified sensw lato,
However, it is exceedingly variable in size, and shape of the basal assemblage,
and already has been subdivided into a number of species and subspecies on
the basis of these. The validity of many of these differentiates must be
questionable as there is often a complete gradation from one to the next or
to Cl. bicornis sensu lato.
Ruedemann (1908, pp. 80-85, 1947, pl. 72, fig. 52) and many others have
shown the variability of the basal assemblage and size of the rhabdosome,
and Fig 1, b-f shows. the great variation found at just one locality. This
makes one wonder at the value of separating the species into a number of
subspecies as there must be only arbitrary divisions between them and they
are apparently of little stratigraphic use (many different forms found
together). So far apparently only three specimens (two of Cl. venustus
and this one) have been found indicating that this stage of development of
Cl. bicornis would be of academic rather than practical interest. The specimen
here described is therefore not separated from, but merely indicated to be,
a Subspecies of Cl. bicornis.
EVOLUTION
Hsu envisaged a line of descent (see Fig. 2, a-d) from Cl. bicornis with
straight extended spines, step (i), through one with drooping spines (ii),
then drooping spines with short projections (iii) to Ol. venustus with large
projections, step (iv). The finding of this specimen would rather indicate
that step (iii) should be a species with drooping spines and only one fully
grown projection, and an extra step with two fully grown extensions added
before the final stage Ol. venustus (h-a). Hsu’s concept of small size and
excess spinosity pointing to the probability of it being a relic form seems
to be backed up stratigraphically here.
230 CLIMACOGRAPTUS BICORNIS WITH A MODIFIED BASAL ASSEMBLAGE
MATERIAL
Only one specimen was found, preserved as a flattened metallic film in
fine black slate. The specimen was incomplete, both the distal and part of
the proximal end lying off the slab (see Fig 1, a). The graptolite had
undergone deformation before fossilisation being bent about 1 cm. from the
sicula and the distal portion undergoing some torsion as jwell. The specimen
was given to the National Museum of Victoria where it holds the number
P26392.
Horizon AND LOCALITY
The specimen came from the Shoalhaven River Gorge near Tallong,
N.S.W., and from Sherrard’s locality 9 in that area (Sherrard, 1949, p. 77).
From this locality Sherrard has identified Dicellograptus angulatus, D.
caduceus, Climacograptus bicornis, Cl. tridentatus, Cl. minimus, Orthograptus
truncatus pauperatus, O. calcaratus basilicus, Cryptigraptus tricornis and
)
[
\
ef f g
h
h — gh
Fig. 2. Postulated evolutionary sequence of forms developing from Cl. bicornis, all
approximately x2. (a) to (d) as proposed by Hsu (1959, pl. 1, figs 7, 11, 12, 13),
(a), (e) to (h) as proposed in this paper.
Retiolites (Plegmatograptus) nebula, from which she had deduced an age of
Elles and Wood’s “Zone 12” or the Dicranograptus clingani zone (op. cit.,
pp. 64, 80, Elles and Wood, 1912). She also further subdivides the zone,
stating that locality 9 represents a higher part of it. This is lower than
Hsu’s Wufengian or Ashgillian (zone of Dicellograptus complanatus) and
would support the supposition that this is a link between Cl. bicornis and
Cl. venustus.
APPENDIX
_ This area (Tallong) is distinctive in that all the species related to Ol.
bicornis have their basal appendages enlarged to greater than normal size.
Sherrard says (op. cit., p. 69) “The development of the appendages in the
former varieties which Ruedemann has studied and figured (1908, p. 80,
Plate A) can be paralleled and surpassed at this place, where the length of
the virgella in Cl. tridentatus and the size of the wings on the shield in Cl.
peltifer greatly exceeds anything shown in Ruedemann’s plate.” Apparently,
HENRY MOORS 231
conditions which suited the function of an elaborate basal appendage must
have prevailed in the environment where these forms lived during this period.
(See Hsu op. cit., p. 351 for suggestions and references.)
Acknowledgements
The author wishes to express his thanks to Dr. F. C. Beavis, Department
of Geology, University of Melbourne, for correcting the manuscript.
References
Butman, O. M. B., 1947.—The Caradoc (Balclatchie) graptolites from limestones in
Laggan Burn, Ayrshire. Part III. Mon. pal. Soc., ci.
Ewes, G. L., and Woop, H. M. R., 1912.—British graptolites. Mon. pal. Soc., \xiv.
Hsu, S. C., 1959.—A newly discovered Climacograptus with a particular basal appendage.
Acta pal. Sinica, Vol. 7, No. 5: 346-352.
RUEDEMANN, R., 1908.—Graptolites .of New York. New York State Museum, Mem. 11.
, 1947.—Graptolites of North America. Geol. Soc. Amer., Mem. 19.
SHERRARD, K. M., 1949——Graptolites from Tallong and the Shoalhaven Gorge, N.S.W.
Proc. Linn. Soc. N.S.W., 74: 62-82.
CHROMOSOME LOCATION AND LINKAGE STUDIES INVOLVING THE
Pm3 LOCUS FOR POWDERY MILDEW RESISTANCE IN WHEAT
R. A. McIntosu and EH. P. BAKER
Department of Agricultural Botany, University of Sydney
(Plates XvVI—xvII)
[Read 31st July, 1968]
Synopsis
The chromosome location of the incompletely dominant gene conditioning resistance
to powdery mildew in Asosan wheat was confirmed as 1A by nullisomic F, analysis.
This gene, formerly designated Ml,, is now known to be an allele at the Pm3 locus and is
redesignated Pm3a following the recommendations for gene nomenclature in wheat.
Repulsion phase linkage studies, using F, genotypic classification verified from progeny
tests, indicated a crossover value of 4:81 + 0-52 per cent between Pm3a and the gene
Hg conditioning pubescent glumes.
INTRODUCTION
The loci for certain of the genes conferring resistance to powdery mildew,
Erysiphe graminis DC. f. sp. tritici Em. Marchal; in common or bread wheat,
Triticum aestivum L. ssp. vulgare (Vill.) have been located on specific chromo-
somes by aneuploid analyses (Sears and Rodenhiser, in Sears (1954) ; Nyquist,
1957; Briggle and Sears, 1966; Law and Wolfe, 1966; McIntosh, Luig and
Baker, 1967). In other instances linkage values have been determined between
genes for mildew resistance and previously localised factors which condition
other characters (e.g., Briggle and Sears, 1966). Such studies on gene location
and linkage intensity have contributed to the rapid expansion of the wheat
genetic map in recent years. The studies to be reported are concerned with
the chromosome location of a gene conferring mildew resistance in the
variety Asosan and with the determination of the linkage value between this
gene and that conditioning pubescent glumes.
LITERATURE REVIEW
Pugsley (1961) identified a dominant gene MI, for seedling resistance
to powdery mildew in the wheat variety Asosan. Briggle (1966) assigned
permanent numerals to three loci conditioning mildew resistance, giving the
symbol Pm5 to the locus at which M1, is located. Briggle and Sears (1966)
reported the presence of a multiple allelic series at this locus in various
wheat varieties. Alternatively they proposed that close linkage could be
implicated. On the basis of allelism they suggested that as many as five
alleles jwere involved. One group of varieties included Asosan, in which the
gene for resistance was effective from the first-leaf stage to maturity. A
second group included the variety Indian 1A, in which resistance was not
expressed until the three to four-leaf stage. In the studies by Briggle and
Sears tests for mildew reactions on the progenies of disomic F2 plants from
PROCEEDINGS OF THH LINNEAN Society or New SourH WALES, VoL. 93, Part 2
R. A. MCINTOSH AND EB. P. BAKER 233
crosses between the Chinese Spring monosomic series and Indian identified
no chromosome unequivocally as the carrier of Pms. However, tests of Indian
substitution line 1A, in which chromosomes 1A of Indian were substituted
for their homologues in Chinese Spring by means of a series of six back-
crosses to Chinese Spring monosomic 1A, showed clearly that this chromo-
some carried Pm3.
A single gene difference between pubescent and glabrous glumes has been
indicated in most instances (see Ausemus et al. (1946)). Sears (1953) by
nullisomic analysis placed: the single dominant gene for pubescent glume Hg
(Tsunewaki, 1966) in the variety Indian on chromosome 1A.
From backcross data Briggle and Sears (1966) found strong linkage in
_ coupling between Hg and Pm3, the crossover value being 0-82 map units.
MATERIALS AND METHODS
A backcross mildew-resistant derivative Asosan x Federation? W2583*
was used in the investigations. It carries the mildew-resistance gene Ml,
from Agosan, possesses glabrous glumes and is distinguished morphologically
from Federation in having grains with red pericarp.
The chromosome location of Ml, was determined by crossing Asosan x
Federation’ as the pollen parent with the series of twenty-one Chinese Spring
monosomics, which are mildew susceptible. Monosomic F, plants were
distinguished from disomic sibs by meiotic examinations of pollen mother
cells, stained with acetocarmine, from anthers in spikes fixed in Farmer’s
fixative. The segregation ratios for reaction types on the primary seedling
leaves were studied in the progenies of monosomic F; plants in each cross.
The mildew susceptible variety Yalta W1373, with pubescent glumes due
to the gene Hg, was crossed with Asosan x Federation® in order to estimate
the recombination value between Ml, and Hg. The progenies of five Fy
plants were studied, about half the spikes on each plant being bagged to
prevent possible outcrossing. Since on previous evidence recombination
between the genes under study was rare, outcrossing could affect estimates
of the recombination value. Seed on bagged spikes of each hybrid plant
was threshed and bulked, but kept separated from seed produced by open
pollination. Each plant progeny was analysed separately. F. seedlings were
classified for mildew reaction type on the primary leaf, sprayed with an
appropriate fungicide to control further mildew development, transplanted
to the field and grown to maturity. The phenotypic classification for glume
pubescence was made macroscopically. In doubtful cases a binocular micro-
scope was used for final classification. Segregates with pubescent glumes
were classified into two classes, one fully pubescent and the other inter-
mediate in degree of development and in length of the trichomes on the
glumes.
The mildew reactions of progenies from F» plants classified for mildew
reaction type and degree of glume pubescence were determined. On the basis
of these tests residue seed from apparent F. recombinants between M1, and
Hg was space-planted in the field and Fs lines classified for pubescent versus
glabrous glumes at maturity.
A strain of wheat powdery mildew designated S.U.1 (McIntosh and
Baker, 1966) was used in the investigations. Mildew reaction types were
scored according to the scheme described by Newton and Cherewick (1947).
* Refers to Sydney University Wheat Accession Register.
234 CHROMOSOME LOCATION AND LINKAGE STUDIES-IN WHEAT
EXPERIMENTAL RESULTS
Chromosome location of Pm3 locus
Asosan X Federation® was virtually immune to mildew, exhibiting “0;”
reaction types on the primary seedling leaf, in contrast to the susceptible
“3+” reaction type pustules shown by the Chinese Spring monosomics.
Asosan x Federation® showed some mildew on the coleoptile, but this was
ignored in the investigations. Hybrid plants exhibited slightly higher (“1”)
reaction types, indicating incomplete dominance of the resistant reaction
type. Segregation ratios for mildew reaction type in populations from mono-
somic F, plants from crosses involving various Chinese Spring monosomics
are presented in Table 1. Some resistant seedlings showed reaction types
similar to Asosan x Federation®, but the majority, presumably heterozygous
in genotype, exhibited “1” types similar to the F; plants. Populations from
TABLE 1
Segregation for seedling mildew reaction type in progenies of monosomic F', plants from crosses between
the various Chinese Spring monosomic lines and Asosan x Federation®
Reaction types
Chromosome ————— Total 13 :1) P value
involved 2 (Qe. bale 34” ;
(resistant) (susceptible)
1A an ee ae 224 15 239 44-69 0:001
1B es a ae 119 40 159 0-002 0:99-0:95
1D a nO) Ay: 51 15 66 0-18 0-95—-0-50
2A (IT) Fs 5s 43 15 78 0-02 0:95-0:50
2B (XIII) 56 ae 60 16 76 0-63 0-50-0-20
2D a ne aie 62 18 80 0-27 0:95-0:50
3A Bas of My: 52 19 71 0-12 0:95-0:50
3B 48 16 64 0-00 1:00
3D 44 19 63 0-89 0:50-0:20
4A 78 27 105 0:03 0:95-0:50
4B 43 20 63 1-53 0-50—-0-20
4D 54 20 74 0-16 0:95-0:50
5A 51 21 72 0:67 0:50-0:20
5B 56 16 72 0:30 ~ 0:95-0:50
5D 17 8 25 0-65 0:50-0:20
6A 69 16 85 1:73 0-20-0-10
6B 33 7 40 1-20 0:50-0:20
6D 65 14 79 2-23 0-20-0-10
TA 49 12 61 0-01 0:95-0-50
7B 38 18 56 1-52 0:50-0:20
7D 44 18 62 0-54 0:50-0:20
Total (excluding 1A) 1,076 355 1,431 0:03 0:95-0:50
20 of the 21 monosomics gave segregation ratios conforming with expectation
for a single incompletely dominant factor pair in Asosan. For the cross
involving monosome 1A (XIV) a highly significant deviation (P < 0-001)
was shown on this hypothesis. The deficiency in the number of susceptible
segregates in the cross involving only 1A implies that the gene conditioning
mildew resistance in Asosan is located on this chromosome.
Three seedlings chimaeric for reaction types (“1—’ and “3+”) closely
approaching the parental types were observed among the progeny of the
monosomic 1A F, plant (Pl. xvr). In all cases the longitudinal division
line between the resistant and susceptible sectors was at, or very close to,
the leaf midrib. Presumably chimaerism resulted from loss in an early
embryonic division of the Asosan 1A chromosome in a seedling monosomic
or monotelosomic for this chromosome. Chimaeric seedlings were grouped
with the resistant class on the basis of this assumed origin.
bo
Ww
Or
R. A. MCINTOSH AND E. P. BAKER
Linkage intensity between Pm3 and Hg
The genotypes of Asosan x Federation® and Yalta can be designated
Pm3Pm8és hghg and pmspms HgHg respectively and linkage was studied there-
fore in the repulsion phase. In mildew resistant F2: segregates two distinct
groups of seedlings were observed. One group exhibited “0;” reaction types
similar to the resistant parent and the other “11+” reaction types. Tests in
F; confirmed the latter class as the heterozygous genotype. Glume pubescence
was also incompletely dominant. The intermediate phenotype in the F,2 group
is shown in Plate XVII together with the fully pubescent and glabrous
elumed phenotypes. Behaviour of F3 again confirmed that the intermediate
class was heterozygous. Behaviour of F3 lines for mildew reaction verified
~ generally the accuracy of F2 genotypic classification for mildew reaction type
and indicated few misclassification errors.
An inspection of F, data, in which the genotypic classification for mildew
reaction type was verified or corrected from progeny tests, indicated certain
recombinant classes resulting from crossing over between Pm3 and Hg in
repulsion in F,; gametogenesis. Individuals in such classes were checked by
F; tests for the correctness or otherwise of F2 genotypic classification at the
Hg locus. Progeny tests for this purpose were of homozygous resistant plants
which were pubescent in F2, of plants heterozygous for mildew reaction type
classified as homozygous dominant or homozygous recessive at the Hg locus,
and of homozygous susceptible plants classified as heterozygous at the Hg
locus. Lines in which plants were scored for pubescent versus glabrous glumes
at maturity confirmed genetic recombination and revealed accurate classifica-
tion of Hghg and hghg F2 genotypes. In 3 cases out of 36, plants classified
as homozygous pubescent in F2 were found to be heterozygous on the basis
of Fs; tests: No significant differences in segregation were found in the
populations derived from bagged versus open-pollinated F, spikes and the
data were pooled for analysis. The F, genotypic totals for mildew reaction
type, confirmed or amended in a few instances on the basis of progeny tests,
were 216 homozygous resistant, 432 heterozygous and 194 homozygous
susceptible. However, 44 seedlings classified for reaction type to mildew
failed to survive transplantation and produced neither adult plants for
pubescent glume classification nor seed for progeny testing. These were not
distributed at random in the three mildew reaction categories, 9 being from
the homozygous resistant, 14 from the heterozygous and 21 from the homo-
zygous susceptible classes. Despite spraying for mildew control, survival
was strongly biassed against susceptible seedlings. The final figures were
adjusted therefore in each mildew reaction genotype. F.2 plants which died
were included to remove the possible effect of differential survival on linkage
estimation. The distribution within glume pubescence genotypes of the plants
which failed to survive was based on that shown for surviving plants in each
category. The adjusted numbers in the different genotypes for mildew reaction
and pubescence genotypes are shown in Table 2.
From maximum likelihood equations, recombination between Pm3 and H g
was calculated to be 4:81 + 0:52 per cent.
DISCUSSION AND CONCLUSIONS
The current investigations using nullisomic analysis demonstrated that
chromosome 1A carried the gene M1, for mildew resistance in Asosan. This
confirmed the findings of Briggle and Sears (1966) who used the chromosome
substitution technique to place an allele in Indian on this chromosome.
Briggle (1966) designated the locus at which M1, is situated as Pm3 and
236 CHROMOSOME LOCATION AND LINKAGE STUDIES IN WHEAT
proposed that alleles at a locus be indicated by lower case letters. On the
basis of this recommendation we propose that the gene in Asosan previously
referred to by Pugsley (1961) as Ml, be designated Pm3a. This symbolism
seems logical and orderly, especially as this gene exhibits the lowest reaction
type on the primary seedling leaf of alleles at this locus.
The postulated origin of chimaeras for mildew reaction in three F»2
seedlings in the progeny of a monosomic 1A F, plant provided additional
evidence that chromosome 1A carried the Pm3 locus. In disomic heterozygotes
it would be necessary to invoke an unusually high rate of mitotic instability
to explain their occurrence. It is highly probable that such seedlings jwere
initially monosomic for chromosome 1A carrying the Pmsa allele and hence
TABLE 2
Numbers of plants in various F, genotypes from five hybrids
between Asosan x Federation? (Pm3aPm3a hghg) and Yalta
(pm3apm3a HgHg)*
Number of
Genotype plants
Pm3aPm3a HgHg .. of ae 2: 1-04
Pm3aPm3a Hghg .. Hie a a 18-75
Pm3aPm3a hghg .. ui oF ae 205-21
Pm3apm3a HgHg .. a3 on ef 14-45
Pm3apm3a Hghg .. Sous se a 403-67
Pm3apm3a hghg .. ale re ae 27-88
pmsapm3a HgHg .. BP Ea 53 195-05
pmsapm3a Hghg .. ea on ais 19-95
pm3apm3a hghg .. ae we a 0-00
Total Ge ne ae Wa 886-00
* Adjusted to include 9 mildew-resistant (Pm3aPm3a)
plants, 14 plants heterozygous for mildew reaction type
(Pm3apm3a) and 21 mildew-susceptible (pm3apm3a) plants
which failed to survive after transplantation.
x5 (1 Pm3aPm3a: 2Pm3apm3a: 1 pm3apm3a)=0-26 ;
P=0-95-0-50.
x3 (1 HgHg: 2 Hghg: 1 hghg)=1-15; P=0-95-0-50.
hemizygous for this gene. A high rate of mitotic instability is more charac-
teristic of monotelocentric than normal chromosomes in Agropyron addition
lines to wheat (Baker, unpublished); Steinitz-Sears (1966) reported also
that telocentrics for the short arm of monosome 3B in wheat were unstable
Somatically. This suggests that the chimaeric seedlings may have been
produced, in fact, from individuals monotelocentric for the arm of 1A carry-
ing the Pm3a gene. Assuming the constitution of the chimaeric sectors is as
postulated, a comparison of the reaction types in the resistant sector with
those exhibited by Asosan x Federation? (Pl. xv1) indicates that Pm3a is
slightly less effective in the hemizygous than the homozygous state. However,
the reaction types appeared lower than in the heterozygous state in identical
backgrounds under the same environmental conditions.
In crosses both with Chinese Spring monosomics and Yalta, Pm3a was
incompletely dominant. The heterozygous genotypes in crosses with Yalta
showed somewhat higher (“11+”) reaction types than with Chinese Spring
(“1”). This may have been due to the higher temperatures prevailing when
the Yalta crosses were tested.
R. A. MCINTOSH AND EB. P. BAKER 237
The estimate of linkage intensity between Pm5 and Hg in coupling by
Briggle and Sears (1966) from testcross data, using the variety Chancellor
with the double recessive genotype as the male parent, indicated close
linkage with a crossover value of 0-82 per cent. The value of 4-81 + 0-52 per
cent in the current investigations from F,2 studies with linkage in repulsion
is significantly higher. In maize there is evidence that crossing over for
many of the chromosomes is considerably higher in male than female gameto-
genesis (Rhoades, 1941; Burnham, 1949). Briggle and Sears ‘estimate
restricted crossing over to female gametogenesis whereas in the current
studies crossing over occurred in both sexes. It is not known how wide-
spread the phenomenon of higher crossing over in the pollen is in higher
plants. Ramage (1960), in fact, found crossing over higher in the female in
barley. In any case estimates of linkage frequently vary in different investi-
gations. In barley, for example, Woodward (1957) and Wells (1958) obtained
recombination percentages of 26-5 and 18-0 respectively between the loci for
rough versus smooth awn (Rr) and long versus short-haired rachilla (Ss).
It is of interest that Briggle (1966) described a group of five mildew
resistant varieties, all with pubescent glumes, in which resistance was not
expressed until the three to five-leaf stage and in which three different alleles
appeared to be involved at the Pm3 locus. Selections have been made in
appropriate Fs lines in the current investigations to isolate Pm3aPm3a HgHg
genotypes. These together with the progeny of the single Pm3aPmsa HgHg
recombinant F2 individual classified in the studies will furnish a useful
genetic stock carrying two dominant closely linked markers which are readily
classifiable genotypically and in which the mildew reaction classification can
be made at the primary leaf stage.
Briggle and Sears (1966) from the phenotype of a plant monotelosomic
for the long arm of an Indian chromosome 1A in a Chinese Spring back-
ground and an analysis of its progeny concluded that the Pm3 and Hg loci
were both on the short arm of this chromosome. The distances of each gene
from the centromere can be determined by using a telocentric for the short
arm of 1A in mapping but as yet this aneuploid stock is not available. In
view of the close proximity of the two loci it will be of interest to determine
if the technique is sufficiently precise to place their order with respect to the
centromere should the genes be situated at some distance from it. Should
the loci be close to the centromere the difference between the recombination |
value obtained in the current investigations and that published by Briggle
and Sears (1966) may be due, in part at least, to differences in frequencies
of crossing over in male and female gametogenesis since Rhoades (1941)
found differences more accentuated in regions near the centromeres in maize.
Acknowledgements
Technical assistance was provided by Mrs. L. Roberts and Mr. J. Green.
Photographic assistance by Mr. D. J. S. Gow is gratefully acknowledged.
References
AUSEMUS, E. R., HArrinetron, J. B., Rerrz, L. P., and WorzeLia, W. W., 1964.—A summary
of genetic studies in hexaploid and tetraploid wheats. J. Amer. Soc. Agron., 38:
1082-1099.
BriceLe, L. W., 1966.—Three loci in wheat involving resistance to HErysiphe graminis
f. sp. tritici. Crop Sci., 6: 461-465.
, and SeArs, E. R., 1966.—Linkage of resistance to HErysiphe graminis f. sp.
tritici (Pm3) and hairy glume (Hg) on chromosome 1A of wheat. Crop Sci., 6:
559-561.
238 CHROMOSOME LOCATION AND LINKAGE STUDIES IN WHEAT
Burnuam, C. R., 1949.—Crossing over differences in micro- and megasporogenesis in
corn. Agron. Absts. Amer. Soc. Agron., p. 2.
Law, C. N., and Wo.trs, M. S., 1966.—Location of genetic factors for mildew resistance
and ear emergence time on chromosome 7B of wheat. Can. J. Genet. Cytol., 8: 462-470.
McIntrosH, R. A., and BAxker, HE. P., 1966.—Differential reactions to three strains of
wheat powdery mildew (Hrysiphe graminis var. tritici). Aust. J. biol. Sci., 19:
767-773.
, Luic, N. H., and Baxrr, EH. P., 1967.—Genetic and cytogenetic studies of stem
rust, leaf rust, and powdery mildew resistances in Hope and related wheat cultivars.
Aust. J. biol. Sci., 20: 1181-1192.
NeEwTon, MARGARET, and CHEREWICK, W. J., 1947.—Hrysiphe graminis in Canada. Can.
dhs, Tenens (Oey) 7459 esa
Nyquist, W. E., 1957.—Monosomiec analysis of stem rust resistance in a common wheat
strain derived from Triticum timopheevi. Agron. J., 49: 222-223.
Puestrey, A. T., 1961—Additional resistance in Triticum vulgare to Hrysiphe graminis
tritici. Aust. J. biol. Sci., 14: 70-75.
RAMAGE, R. T., 1960.—Differential crossing over in male and female gametes. Barley
Newsletter, 3: 34.
RuHoaves, M. M., 1941—Differential rates of crossing over in male and female gametes
of maize. J. Amer. Soc. Agron., 33: 603-615.
Sears, E. R., 1953.—Nullisomic analysis in common wheat. Amer. Naturalist, 87: 245-252.
, 1954.—The aneuploids of common wheat. Missouri Agr. Expt. Sta. Res. Bull.
572. 59 pp.
TSUNEWAKI, K., 1966.—Comparative gene analysis of common wheat and its ancestral
species. III. Glume hairiness. Genetics, 53: 303-311.
We tis, S. A., 1958.—Inheritance of reaction to Ustilago hordei (Pers.) Lagerh. in
cultivated barley. Can. J. Plant Sci., 38: 45-60.
Woopwarp, R. W., 1957.—Linkages in barley. Agron. J., 49: 28-32.
EXPLANATION OF PLATE XVI
Portions of leaves of F, segregates from monosomic F, plants in cross Chinese
Spring monosomic 1A x (Asosan x Federation?) showing reaction types to powdery
mildew. Left—resistant segregate. Middle—chimaeric segregate. Resistant (left) and
susceptible (right) sectors. Right—susceptible segregate. (x 8.)
EXPLANATION OF PLATE XVII
F, segregation for glume pubescence in cross (Asosan x Federation?) x Yalta. Left-——
homozygous pubescent segregate. Middle—heterozygous pubescent segregate. Right—
glabrous segregate. (x 3-75.)
Proce.
LINN
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. Soc. N.S.W., Vol. 93,
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PLATE XVI
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Proc. Linn, Soc, N.S.W., Vol. 9:
)
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PLATE XVII
Theat.
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scent glumes
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Phenoty
ne
THE VEGETATION OF THE BOORABBIN AND LAKE JOHNSTON
AREAS, WESTERN AUSTRALIA
J. S. Brarp
King’s Park and Botanic Garden, Perth
(Communicated by Mr. R. H. Anderson)
(Plates xv1lI—xx11)
[Read 31st July, 1968]
Synopsis
The purpose of this paper is to present two vegetation maps from Western Australia
on the scale of 1: 250,000. The two are contiguous and conform to the standard series
of topographic maps on this scale. The locality is east of the Western Australian wheat-
belt in undeveloped country between Southern Cross and Coolgardie, Lake King and
Norseman. Mapping was done from aerial photography. Methods of the survey and the
classification and terminology adopted are outlined. A mapping notation developed for
the purpose is presented for the first time. Factors of the environment are described and
the mapped communities dealt with in detail as to physiognomy and composition. Three
shrubland and five woodland types are recognized which combine variously into five
vegetation systems, in which the plant communities and their related soils occur in
catenary sequences.
GENERAL DESCRIPTION
This paper relates to the two contiguous map sheets of the 1: 250,000
Series, Boorabbin and Lake Johnston which cover country between 31° and
33° S. latitude, 120° and 121:30° E. longitude. The area concerned lies in
general terms to the south-west of Coolgardie, which is only ten miles off
the Boorabbin sheet at its north-west corner. The area then extends 90
miles westwards from Coolgardie towards Southern Cross, and 140 miles
southwards towards Esperance and the coast, excluding Norseman which,
like Coolgardie, lies just off the margin of the area. Road, rail, water and
telegraph communications between the west coast and the eastern goldfields
traverse the north of the Boorabbin sheet. Similar lines of communication
from Coolgardie to Esperance cross the north-east corner of the Boorabbin |
sheet and continue southwards just east of both map margins for their
entire length. A main road crosses the southern part of the Lake Johnston
Sheet from Daniel to Lake King, a road which is readily traversable and
in good condition. A rather less frequented graded track crosses the northern
part of the same sheet from Norseman to Hyden. Otherwise, as the country
is uninhabited, there are virtually no roads and only a few bush tracks.
The first exploration of this area took place from the south under
J. S. Roe in 1849, when Peak Charles was named after the Governor, Charles
Fitzgerald. In 1863 H. M. Lefroy traversed the northern portion. The
Johnston Lakes were named by a prospector, Frank Hann, in 1901. after
the Surveyor-General, H. F. Johnston. Following the discovery of gold at
Coolgardie in 1892 communications with the coast along the present line
of the railway and water pipeline were established, and in 1893 Holland
pioneered a route from Broome Hill near Katanning to Coolgardie. This
route, “Holland’s Track”, crossed the Boorabbin sheet diagonally from S.W.
to N.E. The northern part of it from Pigeon Holes to Coolgardie has
become a graded road, but the southern portion has been lost.
PROCEEDINGS OF THH LINNEAN Society oF NEw SoutH WALES, VoL. 93, Part 2
240 VEGETATION OF THE BOORABBIN AND LAKE JOHNSTON AREAS
We have little precise record of any botanical collecting in the area.
Diels and Pritzel travelled through by train in 1901 and recorded some
observations. In recent years collecting has been done by such botanists
as C. A. Gardner and R. D. Royce and the plant cover is reasonably well
known in a general way. The area was included in the Coolgardie District
of the Eremaean Botanical Province by Diels (1906), which was maintained
by Gardner and Bennetts (1956). Burbidge (1960), however, has placed
the Coolgardie District in an interzone lying between the Eremaean and
Southwestern Provinces. Gardner’s (1942) map of the vegetation of Western
Australia shows the whole area as Sclerophyllous Woodland but this clearly
is intended as a broad generalization, having regard to the scale of the
map (about 1: 8,000,000). Other vegetation maps which include Western
Australia such as in Williams (1955) are equally generalized, due to their
scale and the absence of detailed field work.
The Boorabbin area was geologically mapped by J. Sofoulis and W. Bock
in 1960, and a geological map at 1: 250,000 with explanatory memoir published
in 1963. The Lake Johnston area has not been geologically examined in
detail, nor mapped. In May 1964 the writer travelled through the Boorabbin
area with J. Sofoulis, who explained the correlations between vegetation
and underlying soil and geology which had been observed in the course
of geological mapping. Botanical specimens were collected and numerous
important plants, especially the trees, identified. In October 1964 the writer, |
assisted by F. Lullfitz, traversed the Lake Johnston area from Lake King
to Norseman with diversions to Hatter’s Hill, Lake Hope and Peak Charles,
and returned from Norseman on the Hyden Road as far as Forrestania
with a diversion to the Bremer Range, then going north to Southern Cross
and east along the main road to do spring collecting in the Boorabbin scrub
heath. Four hundred botanical specimens were collected and subsequently
identified, partly by Mr. C. A. Gardner and partly by the Curator and
staff at the Western Australian Herbarium, whose assistance is gratefully
acknowledged. The object of the field work, however, was not to obtain
comprehensive collections to establish the total floristic composition of
each plant community, desirable though that would be. This would have
required very much more time in the field, and in any case there would
have been no facilities for identifying a much greater number of specimens.
The objective has been ecological, to determine the plant communities
occurring in the area, to map them, and to furnish the essential details of
their composition and physiognomy.
Following the field reconnaissance, the mapping was done from controlled
1-mile to 1-inch aerial photo-mosaics furnished by the Department of Lands
and Surveys. Much time was saved by the kindly loan by the Geological
Survey of the original photographs used by the geological field party in
1960 for the Boorabbin area, since after consultation with the party leader,
Mr. J. Sofoulis, his interpretations could be readily adapted for vegetation
mapping. In this part of the country rock exposures are few and the
underlying strata are buried beneath a thick overburden. The geological
map is thus to a large extent essentially a soil map and was prepared by
interpreting vegetation patterns on the aerial photographs as soil. In the
Boorabbin sheet there will be found to be a general similarity between the
boundaries shown on the vegetation and geological maps. The main differences
consist in the subdivision for vegetation purposes of the “Residual Sandplain”
areas into scrub heath and broombrush thicket, and the inclusion of all
the areas of basic rocks and included granites in the northeastern sector
within a single association of sclerophyll woodlands.
J. S. BEARD 241
The Lake Johnston sheet was mapped purely from original photo-
interpretation by the writer on the scale 1: 62,500. The outlines were reduced
photographically by the Department of Lands and Surveys and prepared
for publication at King’s Park.
Factors of THE HNVIRONMENT
Physiography
The country included in these map sheets is a part of the interior
plateau of Western Australia and lies between 700 and 1500 feet above
sea level. With the exception of the group of the abrupt Fitzgerald Peaks
in the south-east, of which the largest, Peak Charles, reaches a height of
2160 feet, relief is subdued. Ridges are broad and flat and alternate gently
with valleys which are also broad and flat, the difference in level from
ridge to flat being rarely more than 300 feet with slopes falling about 25
feet per mile.
The area is divided into five Vegetation Systems (see page 259),
distinguished as the Boorabbin, Coolgardie, Cave Hill, Bremer Range and
Lake Hope Systems (Fig. 1). Each of these has a characteristic physiography,
geological structure and soil association as well as its characteristic
COOLGARDIE
BOORABBIN SYSTEM
LAKE HOPE SYSTEM
LAKE JOHNSTON
Fig.! VEGETATION SYSTEMS
vegetation pattern. In the Boorabbin System the country consists mainly
of a dissected upland developed on granite with widely spreading sandy
plateau surfaces up to 1500 feet in altitude and valleys down to 1200 feet,
draining mainly to the north. In midslope there may at times be a small
lateritic scarp or “breakaway” and also bare outcrops of granite which are
moderately common. They have a low outline and rarely project much
above the surrounding country. The valleys are relatively narrow and contain
chains of small salt lakes. The Cave Hill System is also a granite upland,
but much more strongly dissected, so that little of the old sandy plateau
remains; it is hilly on a smaller scale and granite outcrops are very abundant,
Except for the Fitzgerald Peaks, however, they rarely form prominent
features. Valleys are broader and there are some larger salt lakes. The
Lake Hope System is again a granitic upland, but developed under a higher
D
242 VEGETATION OF THE BOORABBIN AND LAKE JOHNSTON AREAS
rainfall to a still more advanced stage of dissection. Traces of the sands
and lateritic gravels of the “old plateau” surface on high ground are rare
and granite outcrops quite rare. Solonetzic soils carrying mallee vegetation
have developed over the rising ground, valleys are very broad and flat and
contain large salt lakes. Relief is from 800 to 1100 feet above sea level.
The Coolgardie and Bremer Range Systems are developed upon
greenstones, mainly basic rocks such as ancient basaltic lavas and
metasediments, with included granites. These outcrops tend to occur in
belts trending N.N.W.-S.S.E. and form small hilly ranges with rounded
rocky hills and broad alluvial areas at their foot. Relative elevation in the
Bremer Range is only 300 feet.
The physiography of the whole area of these two maps is dominated
by a main watershed (see Fig. 2) running more or less diagonally across
—_-=—-—-—-—_—-—
_—_
=~
= EN
WZ
D
LAKE JOHN: NGS} 7G
Fig.2 DRAINAGE PATTERNS
the Boorabbin sheet from S.W. to N.E. and at a general height of 1300-1500
feet. A second parallel ridge 1000 to 1200 feet high occurs 40-50 miles to
the S.E., and it is between these two ridges that the extensive and
complicated system of the Johnston Lakes has developed. In the whole
region there are no permanent streams, and indeed there are hardly any
well-defined channels for flood run-off. None the less the country shows a
definite pattern of ridge and valley which must have been excavated by
J. S. BEARD 243
a coherent river system, presumably in earlier times under a higher rainfall
régime. Figure-2 is an attempt to sketch the ancient drainage system by
inserting hypothetical streams in the valley bottoms where today there are
only chains of salt lakes with little or no actual flow from one to the other.
Crests of the interfluves are shown dotted. North of the main ridge the
valleys trend northwards and it is thought that this drainage originally
passed westward to the Swan River. On the east side of the Boorabbin sheet
drainage is eastwards towards Lake Lefroy. Elsewhere, south of the main
ridge, the country slopes generally towards the sea, and it is reasonable
to suppose an original connection with those river channels which still
actually exist on the south coast such as the Oldfield and Lort Rivers.
The Bremer Range forms a ridge following the line of strike of the
_ basic rocks, which is thus at right angles to the main direction of the
ridge and valley system developed upon the granite. However, the ridge
is a low one and appears to have been breached by the ancient river
system.
In the present epoch the process must be one of filling up the ancient
valleys by silt and evaporites derived from nearby hills. Some parts of the
valleys would tend to receive more deposit than others, raising the level
relatively more and leading to ponding elsewhere. Evaporation then leads
to accumulations of salts, vegetation is killed at a certain critical level
and playa lakes come into being. These may be extended later by wind
action and by further salt accumulation.
If it is postulated that in the past a period of higher rainfall is
necessary to account for the development of the present topography, or
more than one such period, then one or more periods of greater aridity are
equally reasonable and indeed are evidenced elsewhere on the continent.
During an arid phase aggradation of valleys might well occur more rapidly
than at present, and have initiated the formation of the salt lakes. It
might be possible by careful study to discover the effects of such climatic
fluctuations upon landform.
Climate*
As most of this country is unsettled, rainfall records are available only
for a few places near the northern boundary. Maps based on these records
and others reasonably close to the area show that the 11 inch isohyet runs
roughly northwest-southeast through the northern part of Lake Johnston,
dividing the area approximately into halves. The lowest rainfall is just
below 10 inches in the far northeast, and the highest is 13 inches, on the
eastern part of the southern border. In about 90% of the area annual
rainfall is between 10 and 12 inches, there being only a comparatively
small area near the southern border with 12 to 13 inches, and a still smaller
area in the north with less than 10 inches.
In the southern part the wettest six monthly period is the same as
in most of the agricultural area, i.e. May to October. However, further north,
and in about four-fifths of the area, the wettest period is March—August.
Almost throughout the area, the average rainfall is higher in March than
in April, and in the north, where the late autumn and winter rains are
comparatively light, this results in the March—August period being wetter
than May—October.
In the far south, monthly averages give the impression of almost uniform
rainfall, as at Salmon Gums, for instance, they exceed one inch in the
period March to October inclusive, and are between three-quarters of an inch
*The data which follow were contributed by the Commonwealth Bureau of
Meteorology by courtesy of the Regional Director, Perth.
244 VEGETATION OF THE BOORABBIN AND LAKE JOHNSTON AREAS
and one inch in each of the remaining four months. However, the comparatively
high averages of these four months are due not to consistent totals of
about three-quarters of an inch to one inch, but to many dry months, a
smaller number of moderately wet months and a few very wet ones.
Table 1 shows rainfall data for Coolgardie, Widgiemooltha and Salmon
Gums.
TaBLeE |
Rainfall data
(Points)
Zz
3
Pal
a) = > CoD) > aD 7 5
; | 2 BR aE g 5 a } ® Total
SeOn IAB Sa neq Hogeorsle mace ae
Coolgardie
Average no 63n) (6598, 2.96 Sol 10S eile 93s 98e 54. 68s GO memOommmEOUT
Median PMP DXi OD oA AEB O29 Oy SG, GEE as Sl Ll
Highest month 72 643 933 527 415 454 469 297 356 364 309 306 350
Lowest month 72 0 0 0 0 1 7 0 3 0 0 0 0
Highest day .. 72 462 533 230 203 167 193 112 186 92 259 154 190
Widgiemooltha
Average 2.65 NATE! 198) S1LOV 104 S116 - 11632593) S98 G7 718s Gem Ogs
Median --965; 938 940° 50 60) 9O0y MOD ipa anil tie o0 pa nosey OF oO
Highest month 65 5431,054 592 569 591 463 326 330 384 393 342 407
Lowest month 65 0 0 0 0 0 0 3 0 0 0 0 0
Highest day .. 65 3451,050 220 273 172 220 150 122 147 198 203 208
Salmon Gums
Average PASS So Oy = TO MOA ese ly ye 4 Gime Sil seal Otel OS) and 76 1,341
Median ne 43) 42 58 87 117 1388 140 124 100 80 60 £49
Highest month 43 553 410 694 416 435 410 333 381 374 326 280 288
Lowest month 43 0 0 0 0 4 45 25 21 12 0 0 3
Highest ‘day: *. 40° 355 323 415 178 170) 132) 95. -153" 268" 1207 138 Sat
Evaporation
Evaporation ranges from 55 inches per year at the southwestern corner
to 92 inches in the far northeast. The rate of evaporation is highest in
January, when totals for the month at these places are 10 inches and 15
inches respectively. They are lowest in July, when the range is from
1:7 to 2:5 inches.
Growing Season
The length of the growing season may be estimated from curves of
“effective” rainfall (based on evaporation) and average rainfall. These
values are applicable to introduced species drawing their moisture supply
from a comparatively shallow depth, and are not necessarily applicable to
native vegetation. However, they enable growing conditions in this area
to be compared with those in the agricultural area and are included for
that reason.
The growing season is longest on the southern border of the region,
where it reaches four months, and is about the same length as in much
of the wheat belt along a line from Kellerberrin to Northampton. In about
one-third of the area the length of the season is between four and three
months, this being equivalent to the season in the marginal wheat belt
country. In a little more than one-third of the area the length is between
three and two months, while in the northeastern quadrant it is less than
two months.
J. S. BEARD 245
Although the length of the season in the southern part of this area
is about the same as in much of the wheat belt, there is a smaller surplus
of rainfall here during the growing season. On the other hand, a small
surplus of rainfall above the average for the months preceding or following
the normal growing season, would cause a greater increase in the length
of the season in this region than in the main wheat belt.
Temperature
In summer, temperature increases northward, the range in February,
the hottest month, being from a mean maximum of 85° in the southeastern
corner to 94° in the northwest corner. In winter, mean minimum temperature
_ is lowest in the southern half of the area, but increases towards the southern
border, and also over the northern half of the area. The range of mean
minimum is from a little below 39° in the coldest part to a little over 41°
in the far northeast.
It has been found that light frosts occur with screen temperatures of
36°. In an area with a mean minimum of 39° it is obvious that frosts will
occur frequently, and some will be severe. Table 2 shows temperature data
for Southern Cross, Coolgardie and Salmon Gums, the climatological stations
closest to the area. The frequency of temperatures below 36° has been
included in this table as an indication of frost frequency.
Wind
One station in this area (Boorabbin), recorded wind for 24 years, and
the 9 a.m. records for this centre show that from November to April winds
are mainly easterlies, E./S.E. being a little more frequent than E./N.E. in
January and February, and the reverse in the other four months. There
are very few winds with a westerly component during the December—March
period, but they become more frequent in April, and from May to August
westerlies predominate, though not to the same extent that the easterlies
did earlier in the year. Winds in excess of 30 knots were recorded only
three times, from directions of W.S.W., W. and W.N.W. Most of the winds
in excess of 20 knots were from the northwest quadrant.
At Norseman, winds from the northeast quadrant predominate from
January to May, with W./N.W. winds becoming more frequent in May, and
predominating from June to September. In October winds become more
variable and although there are still a fair number from W./W.N.W. there
are more from S./S.S.E. and from N.E./N.N.E. In the November and
December southerlies prevail, but are not much more frequent than north-
easterlies.
At Coolgardie the prevailing direction at 9 a.m. is N.E./N. from October
to April, and S.W./W. from May to September. At 3 p.m. N.E./N. winds
prevail from November to March, N./N.W. in April, and S.W./W. from
May to October.
Table 3 shows the diurnal variation of prevailing wind at Southern
Cross. In this table winds from the direction preceding the stroke are more
frequent than from the direction following it.
Human. Influences
Owing to the low rainfall of the Boorabbin and Lake Johnston areas
there has been virtually no agricultural settlement, the only developed section
being confined to approximately 900 acres in the extreme south-east centred
upon Kumarl, which lies a few miles east of the sheet boundary. There
are no pastoral leases at the present time, though two areas of 56,400 and
VEGETATION OF THE BOORABBIN AND LAKE JOHNSTON AREAS
246
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J. 8. BEARD — 247
52,149 acres at the Johnston Lakes were taken up from 1955-1957 with the
intention of utilizing country where the woodland has a saltbush understory.
The leases were abandoned due to lack of water supplies. There are
settlements along the goldfields lines of communication across the north
and east of the Boorabbin sheet, but these are not agricultural and have
effected virtually no clearance of the plant cover. There has been mining
development in the Coolgardie system, but this likewise has effected virtually
no clearance of the natural vegetation. All mines are closed down at the
time of writing. There is no longer any Aboriginal population in the area.
On the other hand, woodcutting has been in progress steadily for many
years to provide mining timber and firewood for the goldfields and water
pumping stations. More or less all the woodland in the eastern half of the
Boorabbin sheet was cut over at one time, working from tram lines which
no longer exist, but are shown on the geological map. Most of the woodland
TABLE 3
Diurnal Variation of Prevailing Wind
Southern Cross
Number Direction
Years Jan. Feb. Mar. April May June July Aug. Sept. Oct. Nov. Dec.
0600 5 SE/S SE/S SEH/S SE/S N/NE N/NW NW/W NW/W W/NWSEH/S' SE/S_ SE/S
0900 5 NE/N NE/N NB/N NE/N NE/N N/NE N/NE W/NWNE/NNE/N NE/E NE/E
1200 5 N/NE N/NENE/N N/NENE/N. N/NENW/W W/NWW/SW W/SW W/SW N/NE
1500 5 SW/W SE/S NE/N NE/N W/SW N/NE W/SW W/NWW/NW W/SW W/SW_ S/SW
The wind data are of interest in view of the attention drawn by Sofoulis (1963) in the geological
memoir to the elongation of the playa lakes in a NNW-SSE direction with an apparent migration
of the lakes to the NNW and piling up of sand on the SSE side. During winter, the rainy season
when the lakes may contain water, westerlies predominate and nearly all the strong winds (> 20
knots) are from the north-west quadrant. This would account for the transport of sand to the
south-east, and it must be assumed that the wave motions set up tend to form sandy beaches
on the SE and to undercut the banks in the NW, even though at first sight the opposite would
seem likely to be the case.
on the remainder of the Boorabbin sheet is now being tapped by motor truck
working from roads running south from the Great Eastern Highway. The
Lake Johnston sheet on the other hand is much more remote and untouched,
and timber getting is confined to a belt about 10 miles wide along the
southern half of the eastern margin of the sheet. Fellings which are very
selective have had the effect of making the woodland more open. Regeneration
seems generally to be satisfactory from seed except in LHucalyptus
salmonophloia stands where it may be very slow to appear.
While human land utilization pressure in this area is minimal, it is
quite another matter with that other anthropogenic factor, the bush fire.
All large contiguous areas of scrub, thicket and mallee show burn patterns
in air photographs, so much so that interpretation is often difficult. The
taller sclerophyll woodlands burn only rarely and with difficulty if at all,
owing to the sparseness of the ground layer and openness of the tree canopy,
but the much denser shrublands burn all too easily. In the south, where
woodland is shorter and denser and merges into mallee, it, too, will burn
readily and there may be difficulty in distinguishing on photographs between
mallee and burnt woodland. We can be certain that fire has occurred since
time immemorial in this vegetation, at least as far back as the earliest
248 VEGETATION OF THE BOORABBIN AND LAKE JOHNSTON AREAS
human occupation, but the probability seems to be that it has become
more frequent since the arrival of Europeans. Burn patterns show recent
fires to have started on roads and tracks, or to have spread from settled
country in the direction of Southern Cross. Sofoulis (1963) has drawn
attention to features detectable in burn patterns in aerial photographs which
he describes as “sandplain lineaments”, and attributes to lines of taller
vegetation representing some source of underground water supply such as
faults and joints in the underlying rock. It is difficult to subscribe to this
view since the patterns radiate fanwise from the source of origin of a fire
and extend along the direction in which the fire has moved most rapidly.
Further, in the sandplain country the underlying rock is deeply buried and
unlikely to affect surface vegetation. However, this is something that requires
further investigation.
DESCRIPTION OF THE VEGETATION
Plant Formations
The Plant Formations occurring in this area are only six in number.
Describing them in the simplest terms only, they are as follows:
1. Scrub Heath: Popularly “sand heath” or “sand plain’, a mixed,
stratified, partly open shrub assemblage with Proteaceae and Myrtaceae
prominent, found on leached sands.
2. Broombush Thicket: Popularly “tamma scrub”, a less diverse single-
layered very dense shrub assemblage consisting mainly of Casuarina, Acacia
and Melaleuca species, found on shallow sandy soil underlain by laterite
ironstone and gravel, or by unweathered granite.
3. Rock Pavement Vegetation consisting of lichen and moss on soil-less
outcrops of granite, with aquatic plants in pools and shrubs in crevices
or occasional soil patches.
4. Mallee: Open to closed eucalypt shrub assemblage with variable
low shrub ground layer, found on leached granite soils in the south of
the area.
5. Sclerophyll Woodland:
(a) Mixed sclerophyll woodland, a medium-tall (40-50’) open eucalypt
woodland occupying eluvial soils in the north of the area.
(0) Salmon Gum woodland, tall (> 60’) open eucalypt woodland
occupying alkaline alluvial soils throughout the area.
6. Halophytes: Communities of succulent or more or less succulent
subshrubs occupying highly saline depressions.
All of these formations are well developed upon the acid rocks which
underlie the greater part of the area. Where there are outcrops of basic
rocks, formations of 5(6) and 6 continue to occur as before on alluvial
soils, 1, 3 and 4 are absent, while 2 and 5(a@) occur with a distinctly different
floristic composition.
It will be observed that two different terms, “scrub” and “thicket”
have been used for shrubland formations, and this demands a word of
explanation. Just as it is useful to distinguish between closed and open
tree-dominated formations as ‘forest” and “woodland” respectively, so it
Seems necessary to distinguish in the same way between closed and open
shrublands as “thicket” and “scrub”. “Mallee”, the eucalypt shrubland
which has physiognomic properties of its own, should strictly be referred
to as “mallee scrub”.
J. S. BEARD 249
Physiognomy _
1. Scrub Heath
It is difficult to describe a characteristic structure for this formation,
since the vegetation is burnt so frequently that a mature structure has
little chance to develop. On regeneration after fire by coppice and seedling
growth, a layer of low shrubs appears, at first open and then more or
less closed. With further growth the naturally taller species begin to
outstrip the smaller, and stratification begins to develop into a lower layer
of small ericoid shrubs with emergent taller species with larger leaves.
In time the upper layer, while remaining irregular due to the varying mature
sizes of its components, may tend to close up and partially suppress the
lower layer. The latter grows from 1-24 ft in height, its components
typically belonging to the Myrtaceae, with leaves leptophyll in Raunkiaer
size. Some of these have a corymbose habit of growth, others are erect, thin
and straggly. The components of the upper layer may reach 15 ft in height,
belong typically to the Proteaceae and have very deeply divided, harsh,
prickly leaves which are difficult to assess for leaf area but would be
mainly microphyll in Raunkiaer size. There is some admixture of Casuarina
from the adjacent thicket formation and low mallees tend to occur where
there is some clay in the subsoil. All components are evergreen, with simple
(though in many cases deeply divided) leaves. There is scope for considerable
further investigation into the life form of scrub heath plants, their
regeneration after fire and other aspects.
The structure is illustrated in the diagrams of Fig. 3, which are drawn
from actual measured strips 50 ft long, 10 ft wide and represent what
appear to be early and later stages of development. In the “early” stage
se Po
(
)
wae an wp Ss Ci
ZiONY Ht e@VNYy
154 feet
/
\
SS
Oe \
Yj
\
i
RA
ag C
We, Be
Fig. 3. Profile diagrams of Scrub Heath. Top—early stage. Bottom—late stage. Both
near 255 mile peg, Hyden—Norseman Road. For key to species see p. 257.
my \\
|
)
|
|
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250 VEGETATION OF THE BOORABBIN AND LAKE JOHNSTON AREAS
the tall, pyramidal Grevillea excelsior shrubs have reached 10 ft-15 ft in
height and a lower layer of 2-8 ft high shrubs has become established,
interspersed with small clumps of sedge, small caespitose aphyllous plants
and prostrate woody plants such as Balaustion pulcherrimum and Borya
nitida. In the “later” stage many of the Grevillea are much taller, and
all of them are beginning to show signs of senility. One in the left centre
of the profile is recorded as stagheaded. Broombrush Acacias and
Melaleucas have grown up to 6 ft in height, and there appears to be invasion
in progress by Callitris preissii, Santalum acuminatum and a Hakea sp. In
the vicinity of this profile but not recorded in it, other such invaders
were observed to include Hakea multilineata, Calothamnus quadrifudus and
Casuarina corniculata. The supposed invaders are all relatively young
plants of species which are known to become much larger. It seems probable
that Grevillea excelsior is a pioneer species in scrub heath following fire
and that a succession can be recognized. The ground layer of the “later”
stage differs in that there are large and well established sedge clumps,
with numerous small ericoid shrubs of 1-14 ft.
2. Broombush Thicket
In its purest form the thicket is a dense, closed, single-layered
community of relatively simple floristic composition. It grades, however,
very gradually into scrub heath where the two adjoin, so that there are
intermediate structures such as. the addition of a ground layer of small
ericoid shrubs. Like the scrub heath the thicket is subject to frequent fires
by which it is destroyed, regenerating from seed. Height of the vegetation
therefore depends largely on the time that has elapsed since the last fire,
up to 12 or 15 ft being possible. Where undershrubs occur they naturally
will flourish mainly in the early stages, becoming suppressed later. The
dominant components of the thicket are virtually confined to different species
of Casuarina, Acacia and Melaleuca, all of which have the same habit of
growth: the stem divides repeatedly into a large number of thin, largely
erect branchlets terminating at the same height and giving a dense, gently
domed crown to the bush. This may be referred to as the “Broom-bush”
habit. It is perhaps analogous to the mallee habit of eucalypts. In Casuarina
leaves are reduced to scales and their function taken over by the thin, terete,
green twigs. The fruits are held on the bush until it dies or is killed by
fire, when seed is liberated. The Acacia species have phyllodes, narrow,
linear and about 2 in. long. The seed is shed but is protected by a hard
seed coat: germination takes place copiously after burning. Melaleucas have
small ericoid leaves and their fruits behave like those of Caswarina.
Two profiles have been measured to illustrate the structure of this
formation and appear in Fig. 4, being an “early” and “late” stage as in the
case of the heath scrub. There is no essential difference except that the
virtually closed canopy formed already at about 4 ft in the “early” stage
has advanced to 8 ft in the “late” stage. Heath components such as Banksia,
Hakea and [sopogon have grown rather less in height and are becoming
Suppressed by the broom-bushes.
3. Rock Pavement Vegetation
The numerous bare outcrops of granite rock, devoid of soil, throughout |
this area, are for the most part unable to support any plant cover except |
aquatics in rock pools and some thin crust of lichen. Accumulations of
Peay matter may occur in hollows, which support low woody plants and |
shrubs.
J. S. BEARD 251
4. Mallee =
Mallee is a shrub-eucalypt formation. Each plant has an underground
rootstock about one cubic foot in size from which arise numerous slender
stems giving a bushy crown similar to the “broom-bush habit”. Mallee is
subject to frequent fires which destroy the top growth, regeneration taking
place from coppice. It is not clear to what extent the mallee habit is
genetically controlled or due to fire. Certainly any small eucalyptus tree
of a species having the power to coppice would automatically assume
feet
ie)
~~
Ca
4 ANE,
if “ai a. x Ng NV We Wi \/# ES yi Sey W VN VG
Cama:
Ty
We
rite WH Vf
We al VY il iV Ani MN
aS ua
Fig. 4. Profile ae of Broombush Thicket. Top—early stage. Bottom—later stage.
Top—285 mile peg, Hyden — Norseman Road. Bottom—233 mile peg. For key to
species see p. 257.
\
OWS
‘ya ae ard
SS
= ———
a mallee habit if frequently burnt, and very many mallee species can also
be found in tree form, either moderately sized or in the small but single-
stemmed tree form known as “marlock” in Western Australia. The structure
of mallee is extremely variable, its height varying with age from the last
fire, and its density and associates varying also. The most typical form
of mallee is a closed community of mallee habit rising from 10-15 ft in
height with an understory of small ericoid shrubs of the genus Melaleuca.
The understory may elsewhere consist of mixed shrubs belonging to the
heath scrub where there is a transition to the latter formation, of salt-bush
under alkaline soil conditions or of hummock-grass on red sand. In this area
the last two types occupy only small patches which are not mappable.
The same species of mallee eucalypts may occur over different understories
and vice-versa, and the stature of the eucalypts may vary without change
of species from true mallee to marlocks and small trees. Profile diagrams
measured in mallee are presented in Figs 5 and 6, the former a low form
feet
15
One * nee
ml fot cent
say \
ii AN ATAY
\H ) ‘ Ne We
Fig. 5. Profile diagram of low Mallee, 254 mile peg, Hyden —- Norseman Road.
Wa e Wt
252 VEGETATION OF THE BOORABBIN AND LAKE JOHNSTON AREAS
and the latter a tall. It is believed that these represent early and late
growth stages. In both cases the mallee forms an almost closed canopy
layer, at 7 ft and 20 ft respectively, with a discontinuous lower layer of
Melaleuca broom-bushes. Occasional small clumps of grass were the only
ground vegetation in both cases.
5. Sclerophyll Woodland
_Like the mallee, the sclerophyll woodland is eucalypt-dominated, there
are several different floristic associations and structural types. In general,
growth consists of a tall but open stand of trees with extremely sparse
undergrowth with the result that the woodland is subject to burning to
only a minor extent. Where fire passes, the trees are killed and do not
coppice but regenerate from seed, an even-aged stand resulting. Study is
required of the extent to which existing stands are even-aged, and of the
regeneration of the various species. If it should prove that fire, even at
feet
4 : 20
arp eat? S
Wy Wy
Fig. 6. Profile diagram of tall Mallee near 260 mile peg, Hyden — Norseman Road. For
key to species see p. 257.
very long intervals, is the agent responsible for regeneration of these
woodlands, then as with the other formations height and density are a
function of age and structure is not altogether meaningful. There is in
general a correlation between height and density in that the lowest woodlands
tend to be the densest, and the tallest the most open. There is every gradation
from the low dense mallee to the very tall stands of over 80 ft in height
where the trees are extremely scattered. However, one may distinguish two
broad classes, the mixed woodlands of 40-60 ft in height on residual soils,
and the Salmon Gum (Hucalyptus salmonophloia) woodlands of alluvial flats,
which exceed 60 ft. In the former type trees are irregularly scattered so
that in part their crowns touch and in part there are large open spaces.
Diameters of the dominants are 9-12 in. The trunk forks into a number
of ascending branches at about a third of tree height and the rather flat
crown is thin and casts little shade as the leaves hang downwards. Except
at times in gaps the only tree layer is that of the dominants forming the
canopy, but there are two highly sparse layers of shrubs, the one 6-12 ft
high and mainly of “broom-bush” habit, the other of low shrubs under 2 ft.
Locally the latter may be saltbush. On soil derived from granite the trees
in this formation are smooth-barked, but on basic rocks the majority of the
species have persistent rough bark on the lower trunk or on the trunk and
lower limbs. The significance of this is not understood. Fig. 7 illustrates
a measured profile in an example of this mixed woodland in which Hucalyptus
J. S. BEARD 253
transcontinentalis is dominant. One outstanding tree reaches 75 ft in height,
but most of the dominants attain between 40 and 60 ft. There is an understory
of eucalypts in mallee form. Low shrub and herb layers are virtually absent.
The profile demonstrates the irregularity of the woodland and the openness
of the canopy. There are wide gaps between groups of dominants which
ro) feet
Et ) SN
40 Si (// |
y
20 A my | cana
ce | ee Naan
Fig. 7. Profile aie kate of sclerophyll woodland —Hucalyptus transcontinentalis. For
key to species see p. 257.
tend to occur in clusters, the result no doubt of group regeneration. Much
more work could usefully be done on the structure and stability of these
woodlands. |
The taller woodland of Hucalyptus salmonophloia and its associates is
generally over 60 ft tall, with a maximum of about 90 ft, very open with
the trees as much as 200 ft apart. There is a very strong trunk, in most
cases up to 3 ft in diameter and extending to half the height of the tree.
The shrub layers are as in the mixed woodland, a saltbush understory being
common in the vicinity of salt lakes and on alluvia derived from basic
rocks. Both smooth and rough-barked trees are present but, as before, the
rough bark appears to indicate a higher base status in the soil.
Fig. 8 shows a profile measured in pure salmon gum woodland, an
example which is probably of a denser stand than normal (compare
O feet
BSS
A ait
60 PVE
; Es
Wee
ee be
(ie (as Pep
40 LX } / Wy
We |
20
“re ; pee a O A Gi) dogs Dif NPN
/ Lk \ i, y Wo CN p
Wwe SP Sonos Ae GAY 9 Gop Paar
Fig. 8. Profile diagram of sclerophyll woodland — Hucalyptus salmonophloia association
measured 26 miles south of Hyden on Hyden—Newdegate Road. For key to species
see p. 257.
254 VEGETATION OF THE BOORABBIN AND LAKE JOHNSTON AREAS
photograph). Density is frequently irregular and clustered as with the
Eucalyptus transcontinentalis woodland, and a typical group occurs in the
centre of the measured profile. Height of the dominants is generally greater
than that of Hucalyptus transcontinentalis and its associates. Seedlings,
saplings and young pole-sized Hucalyptus salmonophloia are a rarity,
but there is very frequently a tall shrub layer of the Melaleuca species known
as “Boree”’.
In all the eucalypt types including mallee, the adult leaves are evergreen,
simple, mesophyll in Raunkiaer size, falcate and pendant, hanging vertically.
There is a complete absence of lianes and epiphytes, and of any such special
plant forms as palms, cycads and bamboos, even grass trees.
6. Halophytes
The most extreme communities of halophytes are the small (< 12 in.)
succulent chenopodiaceous shrubs which occupy raised beds on the floors
of the larger salt lakes. Rather rarely here, one may find a Frankenia zone
of small ericoid shrubs round the lake margin. Alluvial flats bordering lakes
tend to have a saltbush understory in woodland and in some places on the
north-east side of a lake the tree cover may be very sparse or virtually
absent, leaving a pure saltbush community. Sandhills to the south-east of
lakes tend to carry a woodland of special floristic composition which is
described in the appropriate place, but has not been mapped. Halophyte
communities have otherwise been mapped where they occur.
Classification, Terminology and Notation
The broad principles of classification adopted are those stated by the
author in previous work (Beard, 1944, 1955). To recapitulate briefly, the
basic unit is the plant association which is a floristic grouping, being the
largest possible group with consistent dominants, either of the same or closely
allied species. Associations may be divided into minor floristic groups, to
which it was proposed to apply the Clementsian terminology. Also they
may be termed consociations if they are single-dominant communities. The
associations may be grouped together according to their physiognomy
(structure and life-fform) into formations. The formation is thus a
physiognomic group and can be treated without reference to floristics.
A higher grouping of formations into formation-series was proposed by Beard
for tropical America (ibid.) and the applicability of this concept to Australia
is discussed below.
Coming down from principles to practical considerations, there is a
need for some consistent and logical system of classification, some consistent
terminology and a mapping notation, for use in the description of vegetation
and in the cartography. On the other hand it is not possible to predict
accurately in advance of a survey what vegetation units are going to be
found in Western Australia, and how they need to be treated. In a general
way, of course, the vegetation types of the State are known, but it would
be unwise to adopt a rigid classification and terminology in advance. It
would be better to consider this towards the end of a general survey and
to adopt in the meantime a flexible system. It is therefore proposed to
distinguish plant formations by local names, e.g., jarrah forest, mallee,
mulga and pindan, and to relate these in each paper as they occur to
recognized classification systems.
At the highest level of classification we have the system put forward
by Kiichler (1949) and elaborated by Dansereau (1951) which was designed
to be of universal application on a world scale and to facilitate valid
J. 8. BEARD 255
comparisons between vegetation units in different parts of the world. A
recent paper (G. Ross Cochrane, 1963) has applied a slightly modified
Kiichler system to the description of Australian vegetation types on a
continental scale. There have been other cases in different countries where
the Kiichler system has also been applied and it seems desirable that we
should also participate. For the benefit of those interested in comparative
vegetation on a world scale it is proposed to give both Kiichler and
Dansereau notations, and Dansereau diagrams, for our vegetation types in
descriptive memoirs, though in point of fact the value of this where
Australian vegetation is concerned may be questioned. Comparisons are
valueless if not valid, and it may be objected that Australian vegetation
has so many unique features unrepresented elsewhere that comparisons
between it and other world vegetation may not be valid at all. The
fundamental idea behind methods of description and classification designed
to be of universal application is that vegetation is the expression of
environment and that if we can equate vegetation units in different areas we
can be assured that they express (within narrow limits) the same environment.
This, however, may very well be a false assumption when totally different
floras are involved in the comparison and it is strongly suspected that this
point will be proven when more exact physiognomic comparisons between
Australian and other vegetation become possible. At least by stating our
vegetation types in Ktichler and Dansereau terms we are making a contribution
to such study, and it can be done without proclaiming any faith in the
outcome of it.
Both Kichler and Dansereau, followed by Ross Cochrane, have spoken
of their systems as “classifications”, but they are in effect descriptive
notations. A classification would appear rather to be an ordered system
of arrangement of vegetation types linked to a terminology, both of which
serve to emphasize and clarify the relationships and differences between
these types. In this sense, a genuine classification of Australian vegetation
types is of the kind published by Williams (1955), with its physiognomic
key and its terminology based on kinds of forest, woodland, savannah and
so on. There appears to be no generally accepted and satisfactory system
of classification for Australian vegetation, but Williams seems to have
been the closest to approach the goal. His treatment contains some inconsistent
features and may not in the long run be workable without modification,
but it may be accepted provisionally and any difficulties can be discussed
as they arise.
In addition to these arrangements for classification and terminology,
we require for use on maps a notation system similar to that employed by
geologists, e.g., Ts, tertiary sandstone; Ag, Archaean granite. In our case
the notation should be of reasonable brevity and should if possible convey
the diagnostic features of the vegetation which are:
1. Nature and size of the dominant stratum and of other strata if of
diagnostic importance.
2. Dominant or diagnostic plant species.
3. Density of the strata referred to in 1.
It should be possible to arrive at a 3-letter notation to embrace the above.
A mapping notation must above all be brief and long alphabetical formulae
which are involved in Ktichler’s system, and even more so in Dansereau’s,
are too cumbersome. The only Australian community cited by
Dansereau (1951), that of Ceratopetalum australiense, has the formula
_Tteazc.Ltehzb.Etegxi.Ftevzc.Hmevzb. and even Kiichler’s much simpler system
Still produces Tropical Rain Forest = Btml.ejuy.HImp. These could never
256 VEGETATION OF THE BOORABBIN AND LAKE JOHNSTON AREAS
appear on any map and it is unlikely that it was ever intended that they
should, the purpose being symbolic description of vegetation, rather than
mapping. However, it is possible to borrow from Ktichler and Dansereau
in the construction of a simpler system as follows:
1. Physiognomy of Dominant Stratum
T Tall trees > 25m. tall
M Medium trees 10—25m. tall
L Low trees < 10m. tall
Ss Shrubs > 1m. tall
Z Dwarf shrubs < 1m. tall
G Graminoids other than spinifex
H Hummock grass (spinifex)
FE Forbs
X Lichens and mosses (Hepatics)
C Succulents (Chemopods)
Ta Tall deciduous trees
Ma Medium deciduous trees
La Low deciduous trees
2. Floristic
Eucalypt
Acacia
Triodia
Heterogeneous (mixed or other)
Hts oO
(To be differentiated as e;; e2 for individual species. Other
species may be added as required, e.g., m = Melaleuca.)
3. Density
Closed or continuous
Open, not touching; incomplete cover
Scattered groups
Rare but conspicuous
Barren — vegetation largely absent
oSHoro
The actual formulae are to be written with the floristic category first,
e.g., eMc; ali meaning respectively a eucalypt-dominated closed medium
tree community (a eucalypt forest in fact) and an acacia-dominated woodland
(mulga). The formulae are designed in this “triangular” form in order
to be more readily comprehended at a glance. The central capital letter
conveys the most important feature, physiognomy; the left hand one floristics
and the right hand one density. This may be found aesthetically more
pleasing, and more efficient, than the usual formula beginning with a capital
and tailing off into a string of small letters whose order and position may
often be confusing.
The principal, capital-lettered category, is based upon Ktichler’s group
1: Height, and is intended to accord mention primarily to the dominant
stratum, e.g., eMc. If two or more strata are considered co-dominant, their
symbols may be written together, e.g., xSZi. Any diagnostically important
strata may be included in this manner—eLr.aSr.pHi—which is the formula
for sandhill desert, i.e., an open Hummock-grassland of Plectrachne with
seattered acacia shrubs and low eucalypt trees. It is not intended, however,
to deal exhaustively with all synusiae in the manner of Dansereau. This
is a mapping notation which sets out to describe salient features only.
J. S. BEARD 257
The floristic category serves to name the species or genera which are
dominant or diagnostic, but as most West Australian communities are of
simple composition or even single-dominant associations, this category will
also to a large extent convey — from the known morphology of the species —
a life-fform characteristic of the community. In Australia a relatively few
genera dominate and provide a characteristic form for whole communities
in this way — Hucalyptus, Acacia and Triodia (or Plectrachne), rather
more rarely Melaleuca, Casuarina, etc. A category “heterogeneous” is provided
for mixed communities in which no definite dominance asserts itself. The
actual species concerned may be nominated if desirable by subscript numbers.
This category is basically the same in conception as Kiichler’s initial one,
which he writes in capitals, using the classes evergreen broadleaf, deciduous
broadleaf, evergreen needleleaf, etc. These were obviously designed for a
North American environment and represent basic life-forms to a large
extent characteristic of taxonomic groups, e.g., conifers and angiosperms.
In Australia life-form is even more closely related to taxonomic groups and
at a lower level. If a community is said to be eucalyptoid, acacioid or triodioid
a characteristic life-form is at once conveyed to the reader.
The third category “Density” is the same as Ktichler’s Group II.
Kiichler’s Group III is not directly included: the characters which it deals
with will have been already incorporated in our series if important, i.e.,
if they are “salient features’, otherwise they are disregarded. Dansereau
has set up three other categories in his system, Function, Leaf Shape and
Size, Leaf Texture. All of these will be found to be conveyed by our first
cataegory Floristic and are therefore not separately required. There are
presented here first a number of actual profiles of vegetation (Figures
3-8), secondly “Dansereau diagrams” (Figures 9-13) in which these diagrams
are converted into pictograms according to the method of Dansereau (1951)
and thirdly a series of comparative formulae according to the system of
notation proposed above and those of Dansereau (1951) and Kiichler (1949)
in Table 4.
TABLE 4
Plant formations of. the Beard Kichler Dansereau
Boorabbin-Lake Johnston area formula formula formula
Serub heath pie gr EX Le Bsze Fteaxi.Fmeaxc
Broombush thicket ape CSC Bse Fmejaxc
Rock pavement vegetation oo XM Li Mljgki
Mallee so Se as .. eSZe Bsze Fleaxc.Fmeaxc
Sclerophyll woodland ey .. eMi Bunli.szr Tmeaxi.Fmleaxb
Halophytes we Ae ae) XH Oik Fljgki
KEY TO SPECIES APPEARING ON DIAGRAMS
A Acacia; B Banksia elderana; C Callitris preissi; Ca Casuarina
acutiwalvis ; Cs Casuarina campestris; Ce Casuarina corniculata; Er Eremaea
pauciflora; Ef Eucalyptus flocktoniae; Eg Hucalyptus gracilis; Eo Eucalyptus
oleosa var.; Es Eucalyptus salmonophloia; Et Eucalyptus transcontinentalis ;
E Eucalyptus species unidentified; G Goodenia sp.; Ge Grevillea excelsior ;
H Hakea; 1 Isopogon scabriusculus; Ma Melaleuca acuminata; Mc Melaleuca
_cordata; Mp Melaleuca pauperifiora; M Melaleuca sp. unidentified; S
Santalum acuminatum; T Thrytomene; V Verticordia.
E
VEGETATION OF THE BOORABBIN AND LAKE JOHNSTON ARBAS
258
Fig. 9. Dansereau diagram for Scrub Heath.
Fig. 10. Dansereau diagram for Thicket.
Fig. 11. Dansereau diagram for Mallee.
Fig. 12. Dansereau diagram for sclerophyll woodland Eucalyptus transcontinentalis —
EH. flocktoniae association.
salmonophloia
woodland — Hucalyptus
sclerophyll
for
diagram
Fig. 138. Dansereau
Association.
J. S. BEARD 259
VEGETATION SYSTEMS
Beard’s work in Tropical America (1944, 1955) based primary
classification upon floristics: (the plant association), secondary classification
upon structure and lifeform (the plant formation) and a_ tertiary
classification upon habitat (formation-series). At the tertiary level, five
series of related formations could be built up with descending structure
radiating from an optimum, each series within the same general kind of
environment or “essential habitat’, their descent reflecting decreasing
availability of moisture or other comparable factors. Under Australian
conditions, while this approach may still be fruitful for certain purposes
and in a fuller state of our knowledge, it would appear at present that for
vegetation survey a regional approach analogous to the Land System
classification used by the C.S.I.R.O. (e.g., Speck 1960; Mabbutt 1963) would
be more immediately valuable.
A Land System is defined (loc. cit.) as an “area or group of areas
throughout which there is a recurring pattern of topography, soils and
vegetation”, and all three aspects of the landscape, together with geology
and climate, are studied and interwoven into Land Research work. A
vegetation survey, like a topographic, soil and geological survey, studies one
aspect In particular with reference to the others, but the same recurring
patterns will be observed and can be used to characterize unit areas. A
vegetation survey will be expected to discover such patterns which will be
dependent upon topographic and/or soil features, in the same way that it
is already accepted practice for a soil survey to recognize recurrent patterns
of soil types dependent upon topographic and/or geological features and
characterized by distinctive vegetation. Working in East Africa, Milne
(1935) discovered that soil types recurred in a definite position according
to slope and originated the concept of the soil catena. Since that time the
catena with modifications has come into general acceptance in British soil
survey work. The Soil Survey of Scotland uses the Soil Association, a term
originated by J. H. Ellis in Canada in 1932, which is a drainage catena.
The term Soil Catena is used in the Soil Survey of England and Wales
for a “sequence of repetitive soil series which recurs in a manner dependent
upon topographic features” (Clarke, 1937), but it is the practice to speak
of Soil Mosaics where the patterns are not topographically controlled. In
the light of existing C.S.I.R.O. Land Research work in Western Australia,
it can be confidently expected that as the Vegetation Survey proceeds patterns
of vegetation will emerge closely corresponding to soil catenas and mosaics
which will be termed catenary sequences of vegetation and vegetation mosaics.
It may be expected that catenary sequences will predominate on the interior
plateau and mosaics, possibly, on the coastal plains.
It will be desirable to recognize unit areas within which the same
patterns recur, and to find a term for these. Although these are geographical
areas, in a vegetation survey they rank as vegetation units. They are not,
therefore, land systems, though it may logically be expected that if complete
land research comes to be done the vegetation of the area will be found
to be the vegetation of a co-extensive land system. We have in this case had
to search for a term for our own use. The term Soil Association suggests
Plant Association as an equivalent, but this is pre-empted for plant
communities at a lower level of classification corresponding to Soil Type.
“Vegetation System” seems to be the best choice of term. Since we have
already from Diels, Gardner and Bennetts a division of Western Australia
into Botanical Districts and Provinces, it will probably be the most convenient
to regard Vegetation Systems as subdivisions of districts. As the Survey
EE
260 VEGETATION OF THE BOORABBIN AND LAKE JOHNSTON AREAS
builds up, the boundaries of the Districts and Provinces, which are at present
highly generalized, will become more exactly defined, and this itself is a
desirable objective.
The relation between the classification adopted in the Vegetation Survey
of Western Australia for vegetation units and major world systems of
soil classification would then become as follows:
TABLE 5
Unit 1 Unit 2 Unit 3 Unit 4 Unit 5 Unit 6
Vegetation Faciation, Plant Plant Vegetation Botanical Botanical
classification society, association formation system district province
etc. (floristic) (physio-
gnomic)
U.S. Soil Soil phase Soiltypes Soil series — = ae
Survey clas- or variant and soil
sification complex
British Soil Subtype Soil type Soil series Soil Major region —
Survey and soil Association
complex (Scotland)
Catenas and
mosaics
(England)
Russian Soil Soil Soil Soil Elementary Soil Soil
Survey individual variety Complex Landscape Region Province
As indicated in the section on Physiography, it is possible to recognize
five vegetation systems in the area studied.
1. The Boorabbin System
At the highest points and in the centre of the widest interfluves, broad
plateaux of deep yellow sand carry a pure scrub heath association. For
several months in the spring this type is filled with brilliantly flowering
plants of all colours. Down slope it merges very gradually, so that
sharp boundaries can never be drawn, into the broom-bush thicket. There
is at first a general mixture then heath plants are relegated to an
understory and finally disappear. Within these two zones outcrops of
granite occur, with their rock pavement vegetation. Down slope under the
broom-bush scrub the yellow sand becomes shallower and is bottomed by
a lateritic hardpan which eventually comes to the surface often with a
small scarp or “breakaway”. Below this there is an abrupt change to
sclerophyll woodland, at first the mixed association of Hucalyptus
transcontinentalis and Eucalyptus flocktoniae on a relatively shallow red
loam soil overlying granite. Further down in the valley bottom on deep
alluvium there is a gradual change to the taller Hucalyptus salmonophloia
association. The latter species is general throughout on light loam. Hucalyptus
longicornis comes in on heavier soil, Eucalyptus salubris on stiff clay and
Eucalyptus melanoxylon where kunkar is abundant.
2. The Cave Hill System
In the Cave Hill System there is very little scrub heath due to dissection
of the sandy plateau surfaces. The interfluves are mainly characterized by
granite outcrops surrounded by thicket growing on shallow decomposing
rock or residual ironstone. On the lower ground the same woodland types
are found as in the Boorabbin system.
J. S. BEARD 261
3. The Lake Hope System
Here there is a relative scarcity of both scrub heath and thicket with
their associated plateau soils. Instead the uplands are covered with the
mallee formation with its typical sand over clay profile. On lower ground
this merges patchily into very variable mixed woodland, variable both in
structure and composition between the extremes of mallee and_ typical
woodland. There is much less of the Hucalyptus salmonophloia type in this
system and it tends to become restricted to “reefs” or narrow belts mainly
in valley bottoms, but also at times on higher ground.
4. The Coolgardie System
This system is mainly developed in the Eastern Goldfields area to the
north-east, and a relatively small portion of it only is included in the
north-east corner of the Boorabbin sheet. The component communities are
mainly sclerophyll woodland with some mallee and broombush thicket. The
basic rocks or greenstones tend to form small, hilly ranges, in which the
highest and stoniest hills are covered with broom-bush scrub mainly of
Acacia, and the less rocky ridges with sclerophyll woodland of the Hucalyptus
torquata-Eucalyptus le soeufii association. It is possible that the Acacia
scrub maybe associated with rock of a particular lithological type. Woodland
on the lower, less hilly and rocky ground corresponds in structure to the
mixed Hucalyptus transcontinentalis woodland of the three vegetation systems
on acid rocks described above, but is of quite different composition,
characteristic species being Hucalyptus le soeufii, Hucalyptus clelandi,
Eucalyptus campaspe.
The commonest of these are “blackbutts”, that is, there is a persistent
scaly bark on at least the lower part of the trunk, whereas on the acid
rocks components tend to be smooth-barked. On low-lying alluvia_ the
Eucalyptus salmonophloia association is found, with a preponderance of
salt-bush in the understory. Occasionally on the higher ground there may
be sandy plateaux, mainly on included granites, the sand being red in colour.
On deep sand, in small patches, one finds mallee with spinifex, a mallee
form of Hucalyptus oleosa with an understory of Triodia scariosa. Otherwise
the plant cover of these plateaux is broom-bush thicket similar to that on
acid rocks.
3d. The Bremer Range System
While the Bremer Range is similar in geology and topography to
the Coolgardie area its plant communities are entirely different. On the
rocky knolls one finds a broom-bush thicket of Casuarina and a complete
absence of Hucalyptus torquata and Eucalyptus le soeufii. In local patches
the thicket breaks away to a grassland with shrubs. Footslope areas carry
a fine sclerophyll woodland of Hucalyptus dundasti and Eucalyptus longicornis
(both blackbutts), with no other associated trees except that where the
woodland has been destroyed by fire the regeneration in addition to the
above two species contains much Hucalyptus corrugata. The latter species
appears to be relatively shortlived in competition with the others and under
undisturbed conditions appears to be confined to a narrow belt between the
woodland and the thicket. On low alluvial country there is the usual
Eucalyptus salmonophloia association.
BoranicaL PROVINCES AND DISTRICTS
From external evidence it would appear that the Lake Hope System,
in which mallee is the dominant member of the catena, forms part of the
Eyre District of the Southwestern Province, whereas the others belong to
the Coolgardie District of the Southwestern Interzone.
262 VEGETATION OF THE BOORABBIN AND LAKE JOHNSTON AREAS
FLORISTIC COMPOSITION
The purpose of this section is primarily to distinguish floristically
the various plant communities recognized. To do this the names of dominant,
diagnostic and common or conspicuous species have to be listed, but there
is no necessity to aim at a complete list of components. This would only
be the objective if a complete botanical survey were being undertaken. As
it is, the plant lists are being made as complete as possible within the scope
of our knowledge, but it should be understood that they are only collectors’
lists and of necessity incomplete.
1. Scrub Heath
This formation is by far the richest floristically of any of those in the
locality and is also without any definite and consistent dominants so that
it is not at present possible to characterize associations within it. There
is scope for interesting and useful ecological research in studying the
composition of this formation in detail. All that can be said as regards
dominance is that Grevillea excelsior and in the northern part Grevillea ptero-
sperma may assume physical dominance by growing to a much greater height
than other components and may at times be quite abundant. They are very
conspicuous and thus tend to appear typical species although their occurrence
is very irregular. On the other hand it is suspected that Grevillea excelsior
is actually a pioneer species and is relatively short-lived, dying out a certain
number of years after the fire which regenerated it and giving way to slower-
growing, slower-regenerating shrubs. There is an indication of this in the
profile diagrams, as discussed earlier.
Further, Grevillea excelsior is essentially typical of deep yellow sand
and occurs sparingly if at all wherever ironstone gravel predominates. Here
Dryandra or Casuarina replace it. There is no distinct boundary between
the scrub heath on its yellow sand and the broom-bush scrub on its ironstone
gravel, the two soils and the two vegetations merging into one another.
The ecotone is characterized by a broom-bush upper layer with Casuarina,
Acacia and Melaleuca, and a heath lower layer of small, ericoid, mainly
Myrtaceous, shrubs. At an early stage after fire the latter is re-established
and appears to be a similar early stage of pure scrub heath. Later,
however, the broom-bushes grow up and suppress it and give the opposite
impression. The character of the vegetation in the ecotone thus varies
according to time elapsed since the last fire.
This process has been described also for the pure heath scrub when
dealing with physiognomy. In the plant lists which follow, an attempt has
been made to define those species which in matured scrub would form
part of the upper layer, as distinct from those which remain part of the
lower layer.
Mallees are found sporadically in the scrub heath and may often, as in
the case of Hucalyptus burracoppinensis and Eucalyptus leptopoda, be typical
of it and not found to occur in any actual mallee formation.
Composition of the Scrub Heath in the Boorabbin System
Tall emergent shrubs: Grevillea excelsior, G. pterosperma.
Upper layer: Acacia beauwverdiana, A. fragilis, A. hynesiana, A.
resinomarginea, A. rossei, A. sterophylla, A. spp. J.S.B.3353-4, Banksia audaz,
B. elderana, B. sp. inedit. J.S.B.38879, Callitris preissii subsp. verrucosa,
Casuarina acutivalvis, C. campestris, C. corniculata, C. dielsiana, C. helmsii,
Dryandra sp. J.S8.B.3871, Eucalyptus burracoppinensis, E. foecunda, E.
J. S. BEARD 263
incrassata, H. leptopoda, EF. platycorys, Grevillea apiciloba, G. biformis, G.
ceratocarpa, G. didymobotrya, G. hookeriana, G. rufa, Hakea falcata, H.
platysperma, H. roei, Isopogon scabriusculus, Persoonia saundersiana,
Petrophile conifera, P. ericifolia, P. semifurcata.
Lower layer: Adenanthos flavidiflora, Baeckea leptospermoides, Balaustion
pulcherrimum, Boronia ternata, Brachysema chambersii, Burtonia hendersonii,
Calytrix breviseta, Chamelaucium pauciflorum, Conospermum brownii, C.
stoechadis, OC. teretifolia, Cyanostegia microphylla, Dampiera lavandulacea,
D. luteiflora, D. stenostachya, Daviesia croniniana, Hremaea pauciflora,
Erichsenia uncinata, Hriostemon brucei, EH. coccineus, Goodenia pinifolia,
Grevillea haplantha, Halgania tomentosa, Henigema dielsii, Hibbertia stricta,
H. uncinata, Jacksonia hakeoides, Lachnostachys bracteosa, Leptospermum
roei, Leucopogon’ sp. J.S.B.3339, Melaleuca cordata, M. holosericea, M.
subtrigona, Microcorys ericifolia, Micromyrtus racemosa, Phebaliwm
drummondu, Persoonia coriacea, Petrophile circinata, Pityrodia caerulea,
P. lepidota, P. uncinata, Plectrachne rigidissima, Pultenaea george, Stylidium
limbatum, Tetratheca efoliata, Thryptomene kochii, Verticordia chrysantha,
V. insignis, V. pennigera, V. picta, V. pritzelli, V. roei, Waitzia acuminata,
Wehlia thryptomenoides.
Composition of the Scrub Heath in the Cave Hill and Lake Hope Systems:
Tall emergent shrubs: Grevillea excelsior.
Upper layer: Banksia elderiana, B. laevigata, B. media, Callitris preissu
subsp. verrucosa, Calothamnus quadrifidus, Casuarina acutivalvis, C. sp.
C. microstachya, Dryandra erythrocephala, D. sp. J.S.B.38681, Dodonaea
stenozyga, Eucalyptus eremophila, BE. flocktoniae, Grevillea sp. aff. aspara-
goides, G. concinna, G. didymobotrya, G. hookerana, G. imcrassata, G.
integrifolia, G. rufa, G. teretifolia, Hakea falcata, H. multilineata, H. roet,
H. subsulcata, [sopogon axillaris, I. scabriusculus, I. sp. inedit. aff. teretifolius,
Petrophile semifurcata, P. seminuda.
Lower layer (shrubs, subshrubs and herbs): Adenanthos flavidiflora,
Astroloma serratifolium, Baeckea crispiflora, Calythrix aff. decandra, C.
breviseta, Casuarina humilis, Chamelaucium megalopetalum, C. pauciflorum,
C. virgatum, Comesperma drummondii, Cyanostegia angustifolia, Dampiera
juncea, D. lavandulacea, D. wellsiana, Hremaea pauciflora, Grevillea
eryngioides, G. prostrata, Halgania integerrima, Hemigenia eutasxioides,
Isopogon villosus, Leptospermum roei, Leschenaultia expansa, L. sp. inedit.,
J.S.B.3785, Logania tortuosa, L. sp. inedit., J.S.B.3737, Lysinema ciliatum,
Melaleuca cordata, M. subtrigona, Microcorys exserta, Mirbelia spinosa,
Persoonia teretifolia, P. tortifolia, Phebalium sp., Pimelea sulphurea, Pityrodia
axillaris, Olearia ciliata, Oxylobium ciliatum, Stylidium bulbiferum, S.
zeicolor, Verticordia chrysantha, V. insignis, V. mitchelliana, V. picta, V.
roei, Xanthorrhoea nana.
As the plant lists are not complete, much of the difference between the
northern and southern areas may be more apparent than real, and represent
the vagaries of collecting. However, there are certainly some real differences,
which in the present state of our knowledge appear to be the following:
Absence in the south of Grevillea pterosperma, Acacia hynesiana, Acacia
rossei, Banksia audax, Casuarina corniculata, Eucalyptus burracoppinensis,
Eucalyptus leptopoda, Balaustion pulcherrimum, Verticordia pritzelii.
Absence in the north of Banksia laevigata, Banksia media, Isopogon sp.
inedit. aff. teretifolius, Adenanthos flavida, Casuarina humilis, Chamelaucium
megalopetalum, Leschenaultia spp., Xanthorrhoea nana.
264 VEGETATION OF THE BOORABBIN AND LAKE JOHNSTON AREAS
2. Broom-bush Thicket
When of mature structure this formation is a closed community, and
the characteristic species are those of its canopy layer. Under certain
conditions an understory of scrub heath species may be present as previously
described, and these need not be further listed. The broom-bush scrub is
developed on two different substrata, ironstone gravels capping granitic
uplands, and rocky knolls occurring in outcrops of basic rocks. The composition
is slightly different in these two cases.
On granite the principal components are: Acacia fragilis, A. resinomar-
ginea, A. stereophylla, A. sp. J.S.B.3400, A. sp. J.8S.B.3415, A. sp. J.S.B.3774,
A. sp. J.S.B.3775, Casuarina acutivalvis, C. campestris, C. corniculata, C.
dielsiana or var., C. helmsti, Eucalyptus foecunda, E. leptopoda, E. oleosa,
Melaleuca acuminata, Tryptomene appressa, T. kochii.
Not all of these species may be present at any one time, in fact it appears
that three species of Casuarina and two of Acacia would be a normal
association, but such local variations have not yet been studied.
Where this community presents a sharp boundary, as it sometimes does
to granite outcrops or to sclerophyll woodland, one may find along the
margin the small mallee Hucalyptus grossa.
On Mount Day in the Bremer Range, which is an outcrop of fine-grained
basaltic rock forming a low dome of altitude approximately 1500 ft, the
top and slopes (except partially in the steepest parts) are clothed with
a broom-bush scrub. This was 4 ft tall in October, 1964, but dead, fire-killed
relics of 8 ft were present. Composition was observed to be: Casuarina
campestris, v.a.; Calothamnus asper, a.; Eremophila sp. 3841, 0.; Dodonaea
ptarmacifolia, 0.; Acacia sp. 3843, 0.; Melaleuca uncinata, 0.; Cassia
eremophila, 0.; Scaevola oxyclona, 0.; Eucalyptus oleosa, forma, o. (mallee,
up to 10 ft); Triodia scariosa, r.
On the steepest part of the east slope the scrub breaks into grass with
scattered shrubs, the grasses including Stipa juncifolia and Aristida arenaria
with herbs Brunonia australis and Waitzia acuminata and the subshrub
Ptilotus obovatus. The major shrubs are the above Dodonaea, Calothamnus,
Acacia, Melaleuca and Cassia with Pittosporum phillyraeoides. At the foot
of the steep slope the dense scrub is reformed, consisting almost entirely of
Casuarina, C. campestris, C. helmsii and OC. sp. inedit. J.S8.B.3838.
Such scrub knolls with grassy openings are frequent in the Bremer
Range, but composition was not examined elsewhere.
Scrub on basaltic hilltops in the Coolgardie system is rather taller and
consists mainly of Acacia sp. 3377.
3. Rock Pavement Vegetation
The numerous granite outcrops throughout this region consist typically
of low domes of almost bare rock exfoliating in thin sheets. The rock is
not quite bare, but clad sparsely with lichens, while here and there are
depressions and holes filled with water after rain which have a certain
growth of algae and mosses, and patches of soil, perched upon the rock. The
vegetation of these patches varies according to their size and depth. Where
they are thin and small, the typical plant is Borya nitida growing in
herbaceous tussocks. With more soil, certain shrubs characteristic of this
habitat appear, notably MKungea sericea, less typically Calothamnus
quadrifidus, Thryptomene australis, Melaleuca leiocarpa and Dodonaea
attenuata, with sedges, Restionaceae and herbs such as [sotoma petraea.
Large patches of deep soil and boulders will tend to develop shrubs and
J. S. BEARD 265
small trees of Casuarina huegeliana and Acacia spp. Around the outer
edge of the outcrop the soil is at first shallow, though supplied with additional
moisture by run-on. Shallow patches tend to resemble those on the rock
itself, but the shrub Melaleuca elliptica is especially typical of this situation.
Deep soil adjoining rocks carries dense groves of Casuarina huegeliana to
20 ft in height, or of an Acacia (unidentified Ac. 3352?). These usually
merge into adjoining sclerophyll woodland with a belt containing Hucalyptus
loxophleba.
' Peak Charles differs rather markedly from other granite outcrops in
size, height, steepness and other respects, and its vegetation has
correspondingly unique features. The rock rises to 2160 feet above sea
level, which implies about 1500 ft above the surrounding country, and consists
of a pink granite: In many places the slopes are precipitous and bare of
vegetation except lichens, but elsewhere patches of soil cling to the rock
and scrub has developed. The Summit is largely bare, with a few old, gnarled
bushes of Leptospermum sp. inedit. (J.S.B.3821), 2 ft tall, growing in
crevices. The middle slopes bear shrub thickets 4 to 6 ft tall, containing
Leptospermum 3821, Calothamnus quadrifidus and Calothamnus gilesii,
Melaleuca fulgens, Callitris preissii, subsp. verrucosa, Baeckea behrii,
Darwinia sp., Hibbertia mucronata, Labichea lanceolata, Anthocercis
genistoides, Philotheca ericoides, Oxylobium parviflorum.
On the lower slopes these also very largely occur with the addition
of other species such as Grevillea teretifolia, Santalum acuminatum and
Santalum spicatum, Acacia spp., Casuarina humilis. Besides this scrub,
however, there are groves of Casuarina huegeliana, a tree up to 20 ft, or
of mallee with Hucalyptus loxophleba and Hucalyptus eremophila. At the
foot of the rock the tall scrub is joined by Acacia acuminata and Pittosporum
phillyracoides.
Kunzea sericea was not observed on Peak Charles, its niche being
occupied by Melaleuca fulgens, which is characteristic of granite outcrops
in the Esperance area to the south-east. The range of Kunzea sericea is to
the north-west as far as the Darling Range.
4. Mallee
The most consistent and abundant species in the mallee formation of |
this area is Hucalyptus eremophila, a variable species which occurs in a
number of different forms and varieties. Associates are Hucalyptus oleosa
vars., Hucalyptus redunca, Eucalyptus incrassata, Eucalyptus pileata,
BLucalyptus leptophylla, Bucalyptus flocktoniae, Eucalyptus lorophleba. Up.
to four species, of which Hucalyptus eremophila is normally one, tend to
occur together in any one locality. Mallee which has resulted from burnt
woodland may contain or consist of Hucalyptus salubris and Eucalyptus
gracilis.
There are many variations in the ground layer beneath the mallee, from
virtual absence to dense and continuous cover of low shrubs 2 to 4 ft tall.
In the latter case Melaleuca pungens is the usual component. On sandy
soil heath shrubs may be abundant, notably Banksia media, Hakea laurina,
Callitris preissii subsp. verrucosa, Melaleuca cordata, Leptospermum roei,
Calytrix aff. decandra, Leschenaultia sp. inedit., Verticordia mitchelliana.
On heavy soil undergrowth is relatively sparse, but includes a number of
typical shrub species, such as Grevillea huegelii, Grevillea oncogyne,
BHremophila calorhabdos, Eremophila dichroantha, Eremophila ?decipiens,
— Olearia adenolasia, Prostanthera arenicola, and Rulingia craurophylla.
266 VEGETATION OF THE BOORABBIN AND LAKE JOHNSTON AREAS
Mallee communities noted on the traverses were as follows:
Lake King-Norseman Road, at S.W. corner of Lake Johnston sheet —
Bucalyptus eremophila, Eucalyptus flocktoniae, Eucalyptus foecunda,
Eucalyptus oleosa var., Hucalyptus sp. unidentified (J.S.B.3740).
Ditto, vicinity of the 100-Mile Tank—Hucalyptus eremophila, Eucalyptus
flocktomae, Eucalyptus pileata, Eucalyptus sp. unidentified, with
white fruits.
100-Mile Tank to Lake Hope—Eucalyptus eremophila, Eucalyptus
foecunda. Young thickets of Hucalyptus salubris and Eucalyptus
gracilis representing sclerophyll woodland in process of regeneration.
Norseman-Hyden Road, 3815-300 Mile Peg—EHucalyptus eremophila,
Eucalyptus gracilis, Eucalyptus foecunda, Eucalyptus redunca.
South of Woolgangie—Hucalyptus eremophila, Hucalyptus loxophleba,
Eucalyptus oleosa var.
Woolgangie to Bullabulling — Eucalyptus eremophila, Eucalyptus
foecunda, Eucalyptus incrassata.
No mallee has been observed in this area on greenstone.
5. Sclerophyll Woodland
In this formation there are a number of different association which may
be readily recognized.
(a) Eucalyptus transcontinentalis*-Hucalyptus flocktoniae association
on the “granite eluvium” of the geological survey, i.e. red loam developed
im situ on granite. Associated trees are Hucalyptus gracilis (0), Hucalyptus
corrugata (f), Eucalyptus salubris (l.a.), Hucalyptus melanoxylon (1.f.).
Undergrowth may be almost entirely lacking, but the few scattered shrubs
seen include: Alyxia buxifolia, Comesperma spinosum, Daviesia anthoclona,
Dodonaea stenozyga, Bremophila dempsteri, Eremophila saligna, Eremophila
sp. inedit. J.S.B.8825, Grevillea huegelii, Grevillea oncogyne, Melaleuca
pauperiflora, .Melaleuca .pubescens, . Melaleuca . sheathiana, . Santalum
acuminatum, Scaevola spinescens, Westringia rigida.
In places, mainly south of the Johnston Lakes, a low woodland may
be encountered intermediate in structure between mallee and sclerophyll
woodland proper, and composed of Hucalyptus flocktoniae and Hucalyptus
eremophila, mostly with a dense understory of Melaleuca pungens.
(b) Hucalyptus aff. striaticalyx-Eucalyptus leptophylla association on
sand ridges. These tend to occur to the south-east of salt lakes, in curved
lines, conforming to the present lake margin. The ridges are low, well
vegetated, and too limited in extent to be mapped. There are good examples
between the 380 and 370 mile pegs on the Norseman—Hyden Road. The
dominant species is a tree suggesting Hucalyptus striaticalyx, up to 40 ft tall,
with persistent stringy bark on the lower trunk, together with a few salmon
gum, Hucalyptus salmonophloia, and the sand salmon gum Hucalyptus
leptophylla. The latter is only a small tree, but may form pure stands
locally. Undergrowth consists of a fairly dense ground layer (2 ft) of
spinifex Triodia scariosa, and the cyperaceous reed Lepidosperma viscidum.
(c) Eucalyptus torquata-Eucalyptus le soeufii association on rocky
greenstone ridges. This association occurs only in the Coolgardie system
and is of limited extent in this area. Hucalyptus torquata and Eucalyptus
* Wor convenience, since these are important ecotypes readily recognizable in the
field, it is preferred to use ths forms Eucalyptus transcontinentalis Maiden and
Eucalyptus longicornis F. Muell. rather than Hucalyptus oleosa F. Muell. var. glauca
Maiden and var. longicornis. F. Muell.
J. S. BEARD 267
le soeufii are co-dominant, abundant and characteristic. Associated trees are
Eucalyptus corrugata, Eucalyptus clelandu, Hucalyptus campaspe and
Casuarina cristata. There is an open shrub understory, largely of Hremophila
spp. up to 6 ft tall and of “broom-bush” habit, notably Hremophila scoparia,
Eremophila glabra, Eremophila oldfieldii, also Dodonaea, Cassia and Acacia
species, interspersed with glaucous 4 ft shrubs of the “Old Man Saltbush’’,
Atriplex nummularia. Forbs include Ptilotus exaltatus.
- (ad) Hucalyptus le soeufi-Hucalyptus oleosa association on deep soils
developed on the greenstones and included granites. This also is confined
to the Coolgardie system and is of limited extent within the Boorabbin
map sheet. Composition is related both to that of the Hucalyptus
transcontinentalis-Eucalyptus flocktoniae and Hucalyptus torquata-
Eucalyptus le soeufii associations, all eucalypt components of both being
present except for Hucalyptus torquata which is entirely absent, being confined
to rocky ridges. A newcomer is Hucalyptus oleosa var. obtusa. Thus we have:
Eucalyptus le soeufii, v.a.; Hucalyptus oleosa var. obtusa, l.a.; Hucalyptus
transcontinentalis (ELucalyptus oleosa var. glauca), a.; Hucalyptus clelandii,
f.; Hucalyptus corrugata, f.; Hucalyptus campaspe, 1.f.; Eucalyptus flocktoniae,
f.; Eucalyptus gracilis, o.
The understory does not differ significantly from that in the Hucalyptus
transcontinentalis-Eucalyptus flocktoniae association.
(e) Hucalyptus dundasii-Eucalyptus longicornis association on deep soil
over greenstone in the Bremer Range. The woodland shows a co-dominance
of Eucalyptus dundasii and Eucalyptus longicornis (Eucalyptus oleosa var.
longicormis), sometimes in mixture, Sometimes in pure patches. The former
of these is characteristic of greenstone soils further east, around and to the
south of Norseman, and the latter of greenstone soils further west around
Forrestania and in other localities. This woodland appears to be rather
readily destroyed or damaged by fire and in many places there are young
saplings and pole-sized stands consisting largely of Eucalyptus corrugata in
addition to the above two species. Since Hucalyptus corrugata was nowhere
observed in mature stands it may be that it is a relatively short-lived pioneer
species: this merits further investigation. In the transition to Casuarina
scrub on Mt Day there is a narrow band of Hucalyptus celastroides and
Eucalyptus sp. (unidentified). Undershrubs are extremely sparse, but the
following were noted: Hremophila sp. inedit., J.S.B.3825; Hremophila
densifolia; Dodonaea stenozyga; Acacia spp. On the low ground a saltbush
Cratystylis conocephala is common.
(f) Eucalyptus salmonophloia consociation on alluvial soils, derived
from both granite and greenstone. The composition of this association is very
simple. Very commonly and most typically the tree layer consists of nothing
but Hucalyptus salmonophloia and this seems to be associated with a light
loam soil. Changes in soil tend to bring in some admixture, Hucalyptus
longicornis on clay-loam, Hucalyptus salubris on stiff clay. Hucalyptus
melanoxylon appears to indicate high base status, usually the presence of
Kunkar. Under ecotonal conditions appropriate mixtures can be seen, e.g.,
with Hucalyptus transcontinentalis, Eucalyptus flocktoniae and_ their
associates or with Hucalyptus flocktoniae and Hucalyptus eremophila (in
marlock form) where the boundary is with mallee. In these cases, Hucalyptus
salmonophloia being of superior stature, a layered woodland is formed. There
is commonly a tall shrub layer of Melaleuca spp., e.g., Melaleuca pauperiflora,
Melaleuca pubescens, and/or Melaleuca sheathiana. Shrubs are extremely
sparse, but include many species of Hremophila and Acacia, Daidiesia
268 VEGETATION OF THE BOORABBIN AND LAKE JOHNSON AREA
nematophylla and Daviesia anthoclona. Where the soil is somewhat saline,
there is a ground layer solely of saltbush, 18 inches high.
6. Halophytes. The vegetation of saline areas may be divided into two
types, saltbush and samphire. The former may be found on alluvial soils
in the vicinity of salt lakes, while the latter occurs in the lakes themselves,
usually on raised beds forming a marked pattern in the lake. Not all lakes
have these beds. The saltbush type begins as a ground layer under Hucalyptus
salmonophloia, Eucalyptus melanoxylon or Bucalyptus flocktoniae, which
become more and more scattered as salinity increases. On the east side of
some lakes the trees may thin out completely to leave pure saltbush and grass
as at Lake Hope, where the saltbushes Atriplex paludosa and Frankema
interioris and grasses Danthonia setacea, Stipa elegantissima were recorded.
Occasionally at lake margins trees of Hucalyptus kondininensis are found, but
this tree does not seem to be common in the area.
Samphires have not been studied and are assumed to be Arthrocnemum
Spp.
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“Vegetation der Erde VII.” Leipzig.
ELLis, J. H., 1932.—Sct. Agric., 12: 338.
GARDNER, C. A., 1942.—The vegetation of Western Australia with special reference to
climate and soils. J. roy. Soc. W.A., 28: 11-87.
, and BENNETTS, H. W., 1956 —The Toxic Plants of Western Australia. ” Perth.
JUTSON, J. T., 1934. —Physiography of Western Australia. Bull. No. 95, Geol. Survey of
Western ‘Australia.
KUcHLER, A. W., 1949.—A physiognomic classification of vegetation. Am. Ass. Amer.
Geog., 39: 201- 210.
MILNE, G., 1935.—Composite units for measuring of complex soil associations. Trans. 3rd
Int. Congr. Soil Sci., 1: 345. :
Ross COCHRANE, G., 1963.—A physiognomic vegetation map of Australia. J. Hcol., 51:
639-656.
SOUFOULIS, J., 1963.—Explanatory notes on the Boorabbin geological sheet. Geol. Surv.,
W.A.
SpPEcHT, N. H., et al., 1960—DLands and Pastoral Resources of North Kimberley area,
W.A., C.S.I.R.O. Land Research Series No. 4, Meibourne.
STEPHENS, C. G., 1961.—The soil landscapes of Australia. C.S.I.R.O. Aust. Div. Soils.
Soils Publ. No. 18.
WILLIAMS, R. J., 1955.—Vegetation Regions. Atlas of Australian Resources, Department
of National Development, Canberra.
EXPLANATION OF PLATES
PLATE XVIII
Top left. Sandheath east of Hyden, October, 1964. Stage of regeneration with abundant
Grevillea excelsior, lower story of ericoid shrubs, Verticordia roei flowering.
Top right. Sandheath east of Lake King near the rabbit-proof fence, October, 1964.
Intermediate stage, well-developed Proteaceous shrubs (Banksia elderana, Isopogon)
and Casuarina with low ericoid ground layer.
Bottom left. Broombush thicket of Casuarina spp. Near Koorarawalyee, October, 1964.
Bottom right. Grassy opening in thicket, Mount Day in the Bremer Range, October,
1964. Foreground: Ptilotus obovatus with Stipa and Aristida, behind shrubs of
Dodonaea, Calothamnus, etc., and Hucalyptus oleosa.
—_—
—
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Proc, Linn. Soe. NSW, Vol. 03, Part + Meare xxtt
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J. S BEARD 269
i PLATE XIx
Top left. Mallee between 100-mile tank and Lake Hope, October, 1964. Hucalyptus
eremophila, E. gracilis. Melaleuca understory.
Top right. Mixed sclerophyll woodland, between 100-mile tank and Lake Hope, October,
1964. Fairly young stand. Hucalyptus transcontinentalis, EL. salubris, LH. melanoxzylon,
FE. gracilis. “Boree”’ Melaleuca at right.
Bottom left. Hucalyptus flocktoniae and saltbush, open woodland in salt country on the
east side of Lake Hope.
Bottom right. Unlogged stand of salmon gum (Hucalyptus salmonophloia) 6 m. south
of Queen Victoria Rock. ;
PLATE xx
Top left. Sclerophyll woodland of alluvial soils. Stand of Gimlet (Hucalyptus salubris)
7 with understory of “boree”’ Melaleuca and Atriplex. Norseman—Hyden Road.
Top right. Sclerophyll woodland on sand ridges. Hucalyptus sp. aff. striaticalyz,
Triodia scariosa. .
Bottom left. Vegetation of a typical granite outcrop. Queen Victoria Rock, May, 1964.
Kunzea sericea in crevices.
Bottom right. Vegetation on middle slopes of Peak Charles. October, 1964. Calothamnus,
Callitris, Leptospermum, Melaleuca fulgens, Restionaceae.
PLATE XXI
Vegetation map of Boorabbin.
PLATE XXII
Vegetation map of Lake Johnston.
Species designated in mapping: E.8, Hucalyptus salmonophloia; E.9, EH. longicornis:
H.10, H. transcontinentalis; H.11, HE. flocktoniae; H.12, torquata; H.13, le soeufi; H.14,
dundasti; B.15, eremophila.
SIR WILLIAM MACLEAY MEMORIAL LECTURE, 1968
WILDLIFE CONSERVATION
H. J. Frito
Division of Wildlife Research, CSIRO, Canberra, A.C.T.
[Delivered 31st July, 1968]
I. INTRODUCTION
The distinguished career of Sir William Macleay in Australia, from 1859
to 1891, covered the period when the inland was being occupied by settlers
and domestic stock. He was in the forefront of this occupation when, soon
after his arrival, he took up country on the Murrumbidgee River west of ene
present town of Darlington Point.
This country is now in the heart of the Riverina, the district that became
the cradle of the wool industry, the location of many of the famous studs,
the birth-place of many of the bush ballads and the folk-lore of shearers and
jumbuks, shanties and riverboats. Yet, when William Macleay went there,
only eleven years after Sturt had probed his way down the river, this was all
in the future. We are told that as late as 1842 “very little was known of the
country situated on the western side of the main road passing from New South
Wales to Victoria on the lower Murray, Edward, Billabong, Murrumbidgee,
Lachlan and Darling Rivers. The general impression was that all this lower
country to the westward was too dry, too flat and too arid for any purpose,
and the few who travelled over it described it as a miserable, wretched, useless
country” (Fletcher, 1893).
The actual site of William Macleay’s squatterage is not certain. There
were Many runs on the Murrumbidgee and the Macleays held several and, no
doubt, some were registered in the names of financial backers rather than the
man actually on the spot. However, by the end of the nineteeth century
between Hay and Darlington Point the various squatterages, runs and blocks
had consolidated into four stations, “Eli Elwah”, “Burrabogie”, “Toganmain”
and “Kerarbury”. The Macleays had had interests in all except the first,
and Sir William had held “Kerarbury”, part of which had formerly been
known as “Uratta’”, since the fifties. ‘“Kerarbury” has since been subdivided
and, although a Kerarbury Station still exists, the block where the original
homestead is thought to have been is immediately to the west and is known as
“The Homestead”.
To a naturalist, however, of greater concern is not that the old homestead
has disappeared and its location is not certain, but that most of the wildlife
of the region has also disappeared and its original composition even is not
certain.
Il. Smartt MAMMALS
In many years’ research on ve Murrumbidgee plains, apart from rabbits,
domestic stock and feral foxes and cats, the only mammals seen have been
red and grey kangaroos, Megaleia rufa and Macropus giganteus, an occasional
marsupial mouse, Sminthopsis crassicaudata, and a few brush-tailed possums,
Trichosurus vulpecula, wherever there is some timber. There are probably also
PROCEEDINGS OF THH LINNEAN SOCIETY OF NEw SovuTtH WALES, VOL. 93, Part 2
H. J. FRITH 271
a few water rats, Hydromis chrysogaster, somewhere on the river, but I have
not seen them. —
In 1958 Marlow published maps of marsupial distribution based on his
own observations and on museum records. The startling fact of these maps
is that apart from the red kangaroo, grey kangaroo, brush-tailed possum and
fat-tailed marsupial mouse, all of which are common today, there are virtually
no historical records of marsupials in western New South Wales. The records
of native rodents are likewise poor. Hither inland New South Wales was
without small mammals at the beginning of settlement, or there has been a
disastrous decline in most species. The latter is, of course, probably the case.
It is one of the tragedies of the early settlement of this country that the
mammal fauna of the inland was allowed to perish without even having been
described. There seems to have been no organized collecting or recording done
in the Riverina, and many animals that could have been common have
disappeared from the State and in some cases from the whole continent.
We will never know with certainty the original composition of the fauna
of the inland plains, and can only try to build up a picture from the few
records from the surrounding areas and from what little we know of the
biology of some of the animals that have survived elsewhere in the continent.
Most of the information on inland marsupials comes from the results of
an expedition led by William Blandowski in 1856-1857 and from the work of
John Gould in 1839-1840. The former expedition was financed by the Victorian
Government and led by William Blandowski, but Gerard Krefft was the
report writer and published papers on the expedition. Wakefield (1966) has
examined the literary accounts and the surviving records. The expedition was
in the field for a year, and, having reached the Murray River near Echuca,
followed it to about the present site of Mildura, where they camped for
Several months. Excursions were made in several directions, including one of
about 300 miles to the north-east. They found a very rich fauna and collected
several hundred mammals representing over 30 species. Judging from the
numbers collected of some species, they must have been very common. John
Gould collected on the Upper Hunter River and on the Liverpool Plains to the
west of the Divide and north-westward from there.
Although both these areas are some distance from the Murrumbidgee, it
is a fair assumption that species collected both at Liverpool Plains and the
junction of the Murray and Darling Rivers would probably also have occurred
along the Murrumbidgee. Among these are the barred bandicoot, Perameles
fasciata, that has not been recorded in New South Wales since Krefft’s 1857
expedition and is not known to occur anywhere today, and Bettongia
penicillata, that has not been recorded in New South Wales for 50 years and is
now only known with certainty in south-western Australia; the bettong was
also recorded near Gundagai in 1832. The rufous rat-kangaroo, Aepyprymnus
rufescens, was common on the Murray, but is now found nowhere inland; the
brown hare wallaby, Lagorchestes leporides, was also recorded near Booligal
in 1890, but this was the last record for New South Wales, and it is not known
to occur anywhere today; the bridled nail-tail wallaby, Onychogalea frenata,
has not been seen anywhere for 30 years; the mouse, Thetomys gouldii, also
seems to have gone as there has been no information for a long time.
Some species recorded by Krefft, but not by Gould, probably did not reach
as far east as “Kerarbury”, but some certainly did. The rabbit-eared bandicoot,
Macrotis lagotis, had a wide distribution in New South Wales, but the last
record was near Wagga in 1912. The pig-footed bandicoot, Chaeropus
ecaudatus, was known in south-western New South Wales until 1880, but is
FF
272 SIR WILLIAM MACLEAY MEMORIAL LECTURE
now probably extinct. lLesueur’s rat-kangaroo, Bettongia lesueurm, was
abundant in much of inland New South Wales, but is now known for certain
only on islands off the coast of south-west Australia.
Kershaw (1909) recorded the hairy-nosed wombat between Deniliquin and
Billabong Creek, not far south of “Kerarbury”, in 1884, and these were
apparently the only examples ever collected in New South Wales. It was
long believed that they were the same as the South Australian species
Lasiorhinus latifrons, but Crowcroft (1967) has shown them to be of the
southern Queensland species L. gillespiei which is also extinct. It may have
also occurred on the Murrumbidgee.
The native rodent, Leporillus apicalis, which is now probably extinct,
probably extended to the Murrumbidgee, and perhaps also Notomys mitchelli.
There may have been other native rodents in the area, particularly in the
Pseudomys group of genera, as Krefft recorded several at the Murray — Darling
junction and Gould and others collected some of these and other species on
the Liverpool Plains.
Marlow points out that of the 52 species of marsupials that have been
recorded in New South Wales, 42% are presumed extinct or rare. Calaby
(1963) considers that of the 119 marsupials in Australia, five are extinct and
34 are endangered. The position with the native rodents is similar but less
well documented, largely because of taxonomic difficulties. Needless to say,
virtually nothing is known of the biology of any of these animals, and
surviving collections in museums are hopelessly inadequate.
One of the significant things about this disaster is that most of the
animals that have disappeared or are endangered have never been persecuted,
or very little. Their disappearance, often unnoticed, is an incidental and
insidious byproduct of alteration of the habitat by stock.
III. KANGAROOS
While the small marsupials of the inland declined before settlement, the
red kangaroo, Megaleia rufa, for a time prospered. Krefft recorded in 1857:
“The Red Kangaroo (Osphranter rufa) is to all appearance very scare as not
a single specimen was brought in during our stay at Gunbower by the natives.”
Many of the explorers’ journals in New South Wales and elsewhere in the
inland also give the impression that red kangaroos were not numerous, -yet
towards the end of the 19th century organized drives in the Riverina and
elsewhere were accounting for many thousands in quite small areas. By 1957
it was widely held that they were in “plague proportions” in the inland, in
places outnumbered the sheep, and were a major pest. There was accordingly
a great deal of research on this and other kangaroos, and it has been possible
to judge these claims and to learn a lot about the biology of the animals.
Should a determined effort be made to conserve them, the essential biological
data on which to base these efforts are now available.
Much of the Murrumbidgee flood plain is now grassland, with scattered
clumps of boree, pine and black box. The most extensive grasslands are of
Danthonia and Stipa/Danthonia. There is general agreement among plant
ecologists (Beadle, 1948; Moore, 1953a, 6; Williams, 1955) that these are
disclimax communities produced by grazing. Originally the area was mainly
a low shrub woodland of boree and other small trees.
Aerial counts of the density of kangaroo populations between Hay and
Darlington Point have shown that in the period 1960-1963 the wallaby grass
plains supported on the average 4:1 kangaroos to the square mile, but of the
woodlands pine-belah supported only 0-1 and the belah-rosewood 1:5 kangaroos
H. J. FRITH Q2v3
to the square mile. Of the total animals counted, 79% were on the wallaby
grass plains (Frith, 1964; Frith and Calaby, 1969). These results, combined
with the common observation that red kangaroos are usually most numerous
on short green grass and least numerous on long dry herbage, leave little doubt
that the abundance of kangaroos in the area is related to grazing on the
virgin habitats.
Newsome (1965) has reported similar results from central Australia with
land grazed by cattle, and there is further evidence from the north-west corner
of New South Wales that the abundance of red kangaroos can be attributed
to sheep grazing also (Bailey, 1967). Haley (1968) in north-west Australia
has concluded that the abundance of euros, Macropus robustus, also is a
direct result of the impact of sheep on a delicately balanced natural vegeta-
tion; the sheep overgrazed it to the point of their own destruction, but created
ideal euro habitat.
This kind of situation, of course, means that kangaroos pose quite
different conservation problems than do many animals. They are abundant
in a particular stage of botanical succession and, if the composition of the
pasture changes due to increases or decreases in the stocking rate with
domestic stock, the size of the kangaroo populations will fluctuate also. In
the meantime, because of their abundance, they are held to be serious
competitors with stock by many, and several States have allowed uncontrolled
slaughter both by pastoralists and by those who shoot the animals for their
meat and hides. The main features of the biology of the red kangaroo are
now known, and it is possible to be rather more definite about the factors that
control kangaroo populations than it was a few years ago when the con-
troversy about kangaroos in inland New South Wales was at its height.
They are not nearly so wide-ranging as has been thought. In mild
weather no marked animal has been found more than 30 miles from the place
of marking, although in drought some have moved up to 130 miles. The
amount of food eaten is not several times that of a sheep as has been often
claimed; in bulk it is similar. It is not possible, however, to equate sheep
with kangaroos as forage removers unless their food preferences are the same.
A current study shows that, although there is a broad similarity in the plants
eaten, there are significant differences. Many plants, relished by sheep, are
avoided by kangaroos and vice versa. The two animals have specific food
preferences and so are not in complete competition. There are similar results
from central Australia with red kangaroos and cattle (Newsome, 1967).
Mortality in the pouch varies with the climatic conditions. In a favour-
able environment loss of pouch young is about 15%, but in mild drought the
number failing to leave the pouch increases (Frith and Sharman, 1964). A
further adaptation to drought is that many females fail to have a_post-
partum oestrus and cease breeding until rain falls.
In the recent drought breeding of red kangaroos ceased, most of the
pouch young and young at foot perished, and the adults dispersed, fell in
condition and many perished. Yet all the time uncontrolled shooting con-
tinued. It is no wonder the populations declined and kangaroos became
uncommon over very large areas. On the Murrumbidgee between 1960 and
1963 the average density fell from 8-6 to the square mile to 2-7, and on the
Darling River it fell from 6-3 to 2-2 and in both places there were later obvious
declines that were not measured (Frith, 1964). In the north-west corner of
the State the density fell from 6-3 to the square mile to 1:2 between 1964 and
1966, and in central Australia between 1959 and 1966 it fell from about 10-0
to 1:9 animals to the square mile (Newsome, 1966).
274 SIR WILLIAM MACLEAY MEMORIAL LECTURE
Faced with this dramatic decline, New South Wales and Northern Terri-
tory strengthened their administrative machinery to control it, but neither
New South Wales nor any other State at present has the manpower to
institute and conduct an effective management programme. Until this is
done kangaroo conservation must remain a matter of chance.
There is no doubt that red kangaroos can be preserved, as a species, in
reserves. However, the animal is a valuable resource and it seems appropriate
that they should also be conserved on land devoted to pastoral industries.
Since the original settlement the stock numbers in inland New South Wales
have followed the common pattern for arid areas. The numbers of sheep
initially are high, but very quickly decline. In the Western Division, for
example, at the end of last century, there were thirteen million sheep, but
due to degradation of the rangeland the area now can only carry seven million.
However, the biomass of animals is probably still much the same, the missing
six million sheep being in the form of kangaroos which are in themselves a
valuable resource. Kangaroo management depends on the Australian com-
munity and Governments realising that domestic stock are not the only
means of utilizing much of the arid zone. A long-term husbandry based on
both stock and kangaroos will probably be less productive than one based on
stock alone, but certainly more permanent.
IV. WATERFOWL
The Murrumbidgee plains have been an appropriate place to study
another complex problem of wildlife conservation, that of several species of
waterfowl. Although the problems of kangaroo conservation are difficult,
they are not insoluble. The position of waterfowl, however, is more com-
plicated as they demand the management of water conservation schemes with
the needs of waterfowl in mind just as much as other products of more
economic value.
Australian waterfowl were apparently never very numerous, and in the
erratic, semi-arid environment one might assume that the populations would
be quite delicately balanced and not need a great deal of interference with
their habitat to upset them. This interference has been applied at several
critical points in the life history of many species and they now are declining
rapidly. The reasons are quite well known, but those responsible for wildlife
conservation have been able to do little to halt the decline.
The inland waterfowl need a breeding place, a nearby refuge for dry
weather and a permanent refuge for extended dry periods and droughts. These
can be separated by great distances, but, unless they are available at the
appropriate time, the populations cannot survive. At present, all these types
of habitat are vulnerable to settlement and are being steadily destroyed.
The inland rivers wind across their flood plains and local rainfall has
little effect on their level; the level is controlled by rain or melting snow on
the highlands, hundreds of miles to the east. In years of adequate rain the
rivers rise, often in spring, but at whatever season the rain falls the billabongs,
lagoons and effluent streams fill. In times of heavy rain there are great floods
that cover thousands of square miles with water a few inches deep. In
extreme years the waters of the Murrumbidgee and the Murray, over one
hundred miles to the south, join across the plains. In dry years the billabongs
do not fill or rise in level only slightly. In very dry years the rivers cease to
flow or become dry and so do the billabongs
In the northern hemisphere, where much of the pioneer work on ihe factors
controlling the breeding seasons of birds was done, birds tend to have fairly
H, J. FRITH 275
regular breeding seasons and nest at a fixed time each year. It has been
concluded that, in general, the sexual cycle is controlled by the increase in
day length that occurs at a fixed time each spring and that the actual breeding
season is timed, within these quite narrow limits, by more immediate environ-
mental factors such as the availability of nest sites, ete. (Lack, 1954; Rowan,
1926). Such a mechanism in semi-arid Australia would, more often than not,
result in the ducklings being hatched at a time when the rivers were low and
the billabongs dry with a resultant disaster to the young of that year.
It has now been shown that Australian waterfowl can breed at any time
of the year that suitable conditions exist for the survival of the young. In
unfavourable years they do not breed at all. The breeding season is directly
_ geared to changes in waterlevel in the billabongs. Even though the plains may
be gripped by drought, should rain on the catchments cause a local increase of
a suitable speed and extent in the waterlevel of the billabong, the birds breed
(Frith, 1959c). In this way it is ensured that the waterfowl do not breed
when the habitat is unfavourable and also that favourable circumstances are
not missed, no matter at what time of the year they occur.
All the common waterfowl of the billabongs receive a sexual stimulus
from the rising waterlevel, but the speed of the reaction varies from species
to species. The grey teal, Anas gibberifrons, is the most rapid, and ovulation
can follow within a few days of the first sign of the rising water. The pink-
eared duck, Malacorhynchus membranaceus, also receives an immediate
stimulus, but the eggs are not laid until it is certain that the rise in waterlevel
is going to be maintained and actual flooding of the plains will follow. This
difference is related to the food requirements of the two birds; the grey teal
can use a very wide variety of food from many sources and this is made
available by the rising water. The most important insects also have breeding
seasons geared to the rising waterlevel and the shallows become dense with
their young by the time the ducklings hatch. The pink-eared duck is a food
specialist and lives on microscopic plants and animals that are only abundant
in shallow drying floodwater. Other ducks have different speeds of response;
the black duck, A. superciliosa, breeds as the waterlevel approaches its
maximum, the hardhead, Aythya australis, breeds immediately after the
maximum level, and the freckled duck, Stictonetta naevosa, a little after that
(Frith, 1959c, 1965). There are two species, the blue-billed duck, Oxyura
australis, and the musk duck, Biziwra lobata, that are restricted to the few
deep semi-permanent cumbungi swamps in the region and, being in a “safe”
habitat, have no need for an erratic breeding season. They breed at a regular
time each year and there is reason to believe that these have retained a
photoperiodic effect in the timing of their breeding seasons (Braithwaite and
Frith, 1969).
How the rising waterlevel stimulates the sexual cycle of species with
erratic breeding seasons has been the cause of some speculation — did the
waterlevel act as a visual stimulus or did it operate through some other
intermediate factor. Recent work with nine species, ranging from those with
the most regular breeding seasons to those with the most erratic, has suggested
that in some species the waterlevel change affects the abundance of food which
in turn affects the nutrition of the birds and this permits breeding. In those
Species having erratic breeding seasons the normal germinal cell division for
Spermatogenesis and the necrosis of these cells are not separated; they occur
concurrently, so that at any time the testis is primed, as it were, for an
immediate response to suitable nutrition of the bird. Both processes are
reversible, so that, should a bird receive a stimulus that is not maintained, it
276 SIR WILLIAM MACLEAY MEMORIAL LECTURE
can rapidly reverse the spermatogenic cycle. Some species can continue sperm
production throughout the year, even during moult (Braithwaite, 1969;
Braithwaite and Frith, 1969).
Floods and even full billabongs are not permanent features of the inland
plains, and as the water areas decrease the birds must disperse. It has been
shown that these movements are multidirectional (Frith, 1959d). The move-
ment is an explosive random dispersal with birds moving in all directions.
The degree of mobility varies from species to species and the distance travelled
depends on the abundance of habitat. The birds apparently fly in straight
lines until a suitable swamp is found and there they remain until it in turn
becomes unsuitable. There is then another random dispersal. A bird having
left the Murrumbidgee only returns by chance. Those that do not find a
suitable swamp on the outward movement carry on, and many presumably fly
out to sea and perish (Frith, 1962, 1963).
Very few of the drought refuges are in the inland as little permanent
water exists there. The few cumbungi swamps that do exist are crowded
each summer with great numbers of many species of waterfowl. It has been
possible to show that, although as many as nine species can crowd together
on the one swamp for long periods, they are effectively isolated ecologically
by having different bill structures, enabling different feeding methods, different
feeding places; e.g., some dive to the bottom of deep water and others feed
only on the surface and have different abilities to use different food items
(Frith, Braithwaite, and McKean 1969). They seldom compete for food. The
cumbungi swamps are capable of supporting the birds for long periods, but
eventually they must move again to the ultimate drought refuges, the coastal
lagoons of the eastern, south-eastern and northern coasts, where they remain
until the inland rivers flood once more.
The causes of the decline in waterfowl populations are now apparent.
The needs of water conservation and hydro-electric power have led to con-
tinuing efforts to control the flow of the inland rivers and to prevent their
flooding. Each water conservation structure built in the highlands and each
weir built on the plains to divert water for irrigation decrease the frequency
with which the water level in the billabongs increases in level and thus
decrease the frequency of breeding of the waterfowl. Even those birds that
do breed are finding it increasingly difficult to find a suitable drought refuge.
The trend of agriculture on the coast is to drain every swamp and pool
whether this will produce productive land or not. Perhaps the productivity
of many farms would be increased more by pasture improvement on existing
pastures than by draining a few acres to produce another boggy tussocky
pasture.
If waterfowl are to survive in south-eastern Australia, it must be recog-
nized that they need living space, and very positive efforts need to be made
to ensure their conservation.
V. WILDLIFE CONSERVATION
Australia’s record towards its wildlife has been generally poor. Those
animals that do not provide products whose value can be measured in money,
or that are not useful as game, have been ignored so that their survival in
the face of increasing pastoral and agricultural use of the land has been
purely a matter of chance. Those that were hardy enough or adaptable enough
to withstand the changes to the environment have survived; those that were
not have declined seriously or have disappeared. Most of those that do
provide economic products have been exploited to the point where they are
incapable of providing anything.
H. J. FRITH 207
I can find no example of successful management of an animal that has
been used to provide economic products. In each case the treatment has
followed the same pattern. The exploitation of the animal is uncontrolled
until the populations have been reduced to a low level. Attempts to control
the industry are then applied, but are often based on legislation rather than
on biology or are not effectively policed or both. The animals continue to
decline in numbers to the point where the industry is no longer profitable,
baving destroyed its own resources. By this time public opinion is so
outraged that the animal is afforded complete and virtually unalterable
protection.
Apart from the few mammals and birds that have been the subject of
special studies, conservation is hindered because the wildlife generally is not
_ well known, so that it is difficult to set targets, and little is known of the
biology of even common species so that reserves can be sensibly managed.
The need for active management of the wildlife in reserves is shown
by. many examples in other countries, but in Australia little has been
documented. There are many cases of reserves failing to serve their main
purpose in the long term because of inadequate knowledge of the animals and
lack of management. I have already mentioned how red kangaroos depend on
short green pastures that is not a permanent feature of the ungrazed inland
plains. It might be inferred that large populations of the animals can only
be retained in reserves if the ground cover is kept in a suitable state by
grazing some domestic stock. On the other hand, the mallee fowl was long
thought to be endangered by the depredations of foxes, but a study of the bird
showed that this was not so; the real reason for the decline was competition
with the birds for food by stock and rabbits; mallee fowl cannot be conserved
unless grazing stock are excluded and rabbits eradicated from the reserve.
The re-discovered Leadbeater’s possum needs an early stage of the regenera-
tion of Hucalyptus regnans with trees 25 to 30 years old so that a dense tangle
of Acacia grows below (Warneke, pers. comm.). Its retention in numbers
would depend on rotational clearing of parts of the reserve.
Colonies of koalas appear on the closely settled coast and often are too
closely protected from interference by the local communities. The problem
with many of these is that they are in small isolated areas of habitat where
regeneration of the trees is prevented by dairy cows and, due to the very
close protection, the koalas increase in numbers to the point where they over-
graze and destroy the mature trees with disaster to the whole colony.
The recently re-discovered New Holland mouse, Pseudomys novae-
hollandiae, promises to show the full cycle of man’s unconscious management
of an area. Only one had been seen in 130 years when it was found to be
abundant near Port Stephens, N.S.W. The reason for its abundance is con-
sidered to be a disclimax community of Acacia and bracken caused by
frequent accidental burning of the forest, but the next phase of the develop-
ment is to destroy the area during mining for rutile and other minerals. But
for the chance encounter by an experienced observer the animal might have
disappeared unnoticed for another 130 years (Keith and Calaby, 1968).
Even if the naturalists of the early days had recognized the extreme
vulnerability of the small ground-living mammals, it is difficult to imagine
how they might have prevented the decline then when we still fail today.
Even had reserves been created on the plains and fenced against stock in an
age when it was not normal to fence even pastoral holdings, this could hardly
have been done before the rabbit invasion which would have destroyed the
habitat anyway. In that age, before it was realized that a knowledge of an
278 SIR WILLIAM MACLEAY MEMORIAL LECTURE
animal’s ecology is essential for its conservation and the management of
reserves, before it was even realized that the science of ecology existed, the
management of any reserves secured on the plains would have been a very
chancy affair. Perhaps the best contribution the early naturalists could have
made would have been to ensure that there was documentation of the animals’
distribution, adequate collections in the museums, and the establishment of
captive colonies against the day when the Governments would be able to
secure and staff adequate reserves.
Today there is improved legislation and a wider interest in conservation.
There are many reserves of varying value and, although few have been chosen
for their wildlife values, this is being rectified in places. Nevertheless, there
are very few reserves where the wildlife is adequately studied and properly
managed so that it will remain abundant. There is no educational establish-
ment that provides training in the techniques of wildlife management.
VI. ConcLUSION
In this talk I have tried to show that wildlife conservation is a complex
process. A great deal of the wildlife has disappeared and many species are
still declining, even though the essential facts to halt their decline are now
known.
The mammal fauna of inland New South Wales disappeared very early
in the history of settlement due to changes in the environment induced by
settlement. It is important to realize that these and similar changes are still
going on, not only in the closely settled areas but the remote ones and in the
reserves, and that, unless care is taken, other wildlife species will fail.
If wildlife conservation in Australia is to be a fact and not merely a
pious hope, a greatly increased effort in research and management is needed.
References
BAILEY, P., 1967.—The ecology of the red kangaroo, Megaleia rufa (Desmarest), in north-
western New South Wales. M.Se. Thesis, University of Sydney.
BEADLE, N., 1948.—‘“‘The vegetation and pastures of western New South Wales.” 281 pp.
Govt. Printer, Sydney.
BRAITHWAITE, L. W., 1969.—Ecology of waterfowl in an inland swamp. V. Testis cycles.
CSIRO Wildl. Res. (in press).
, and Fritu, H. J., 1969—Hcology of waterfowl in an inland swamp. IV.
Breeding. CSIRO Wildl. Res. (in press).
CALABY, J. H., 1963.—Australia’s threatened mammals. Wildlife, 1: 15-18.
Crowcrort, P., 1967.—Studies on the hairy-nosed wombat, Lasiorhinus latifrons (Owen,
1845). 1. Measurements and taxonomy. Rec. S.A. Mus., 15: 383-398. -
HALEY, E. H. M., 1967.—Ecology of the euro, Macropus robustus (Gould), in north-
western Australia. I-IV. CSIRO Wildl. Res., 12: 9-80.
FLETCHER, J. J., 1893.—‘‘Macleay Memorial Volume”, pp. VII-LI. Linnean Society,
N.S.W.
FRITH, H. J., 1959.—The ecology of wild ducks in inland New South Wales. IV. CSIRO
Wildl. Res., 4: 156-181.
, 1962.—Conservation of the Mallee Fowl, Leipoa ocellata Gould (Megapodiidae).
CSIRO Wildl. Res., 7: 33-49.
, 1962.—Movements of the grey teal, Anas gibberifrons Miiller (Anatidae).
CSIRO Wildl. Res., 7: 50-70.
, 1963.—Movements and mortality rate of black duck and grey teal in south-
eastern Australia. CSIRO Wildl. Res., 8: 119-131.
, 1964.—Mobility of the red kangaroo, Megaleia rufa. CSIRO Wildl. Res., 9:
1-19.
, 1965.—Ecology of the Freckled Duck, Stictonetta naevosa (Gould). CSIRO
Wildl. Res., 10: 125-139. 2
, and SHARMAN, G. B., 1964.—Breeding in wild populations of the red kangaroo,
Megaleia rufa. CSIRO Wildl. Res., 9: 86-114.
H. J. FRITH 279
———, and Carasy, J. H., 1969.—“The Kangaroos.” 328 pp. F. W. Cheshire,
Melbourne. _
, BRAITHWAITE, L. W., and McKean, J. L., 1969.—Ecology of waterfowl in an
inland swamp. III. Food. CSIRO Wildl. Res. (in press).
KersuHaw, J. A., 1909—Notes on the hairy-nosed wombat, Phascolomys latifrons, Owen.
Victorian Nat., 26: 118-119.
KeITH, K., and Cauasy, J. H., 1968—The New Holland mouse, Pseudomys novae-
hollandiae (Waterhouse) in the Port Stephens district, N.'S.W. CSIRO Wildl. Res.
(in press).
Lack, D., 1954.—“The Natural Regulation of Animal Numbers.” 343 pp. . Clarendon
_ Press, Oxford.
Martow, B. J., 1958.—A survey of the marsupials of New South Wales. CSIRO Wildl.
Res., 3: 71-114.
Moors, C. W. E., 1953a.—The vegetation of south-eastern Riverina, New South Wales.
I. The climax communities. Aust. J. Bot., 1: 485-547.
, 1953b.—The vegetation of south-eastern Riverina, New South Wales. II. The
disclimax communities. Aust. J. Bot., 1: 548-567.
Newsome, A. E., 1965.—The abundance of red Kangaroos, Megaleia rufa (Desmarest), in
central Australia. Aust. J. Zool., 13: 269-87.
1966.—The influence of food on breeding in the red kangaroo in central
Australia. CSIRO Wildl. Res., 11: 187-196.
, 1967—The distribution of red kangaroos, Megaleia rufa (Desmarest),
about sources of persistent food and water in central Australia. Aust. J. Zool., 13:
289-99.
Rowan, W., 1926.—On photoperiodism, reproductive periodicity, and the annual migra-
tions of birds and certain fishes. Proc. Boston Soc. nat. Hist., 38: 147-89.
WAKEFIELD, N. A., 1966—Mammals of the Blandowski expedition to north-western
Victoria, 1856-57. Proc. r. Soc. Vict., 79: 371-391.
WILLIAMS, O. B., 1955.—Studies in the ecology of the riverine plain. I. The gilgai
microrelief and associated flora. Aust. J. Bot., 3: 99-112.
AUSTRALASIAN MEDICAL PUBLISHING CO. LTD.
71-79 ARUNDEL ST., GLEBE, SYDNEY, N.S.W., 2037
A STUDY OF SOME SMUTS OF ECHINOCHLOA SPP.
_ RR. A. Futuertron and R. F. N. Lanepon
Department of Botany, University of Queensland
(Plates xxIII-xxvitIt)
[Read 25th September, 1968]
Synopsis
The structure and development of sori of Ustilago tricophora on Echinochloa spp.
have been described. There is no essential difference between the sori formed in floral
parts and those occurring in vegetative parts of the host. Sporogenous hyphae growing
_ from the region of the columella form spores progressively, the first formed spores being
those just beneath the soral covering. The latter is composed of a fungal sheath overlain
by a hispid covering of host origin. A study of the development of the smut in both
inocuiated plants and naturally infected plants and a scrutiny of herbarium specimens
of smut of Hchinochloa from many parts of the world have shown that the smut which
occurs in floral organs or in vegetative parts and which is ornamented by spines is
Ustilago tricophora (Link) Kunze. The names Ustilago sphaerogena, U. crus-galli,
U. globigena and U. panici-frumentacei have been applied to smuts which have now been
shown to be indistinguishable from U. tricophora.
INTRODUCTION
Smut fungi are known as parasites of species of Hchinochloa (Gramineae)
in many parts of the world. Seven species of Ustilago have been described
from these grasses, the earliest being an ovary smut, Ustilago tricophora
(Link) Kunze on Hchinochloa colonum from Egypt. Apart from its formal
description by Kunze (1830) almost nothing has been written about that
smut. Another ovary smut, Ustilago sphaerogena Burrill, described in 1888,
has been studied briefly by Fischer (1953) and is recorded from New South
Wales (Anon., 1958). Ustilago crus-galli Tracy & Earle was described in
1895 as affecting both the inflorescence and the vegetative parts, and
McAlpine (1910) has recorded the occurrence of this smut in Australia.
Ustilago panici-frumentacei Bref., U. globigena Speg., U. paradoxa Syd. &
Butl. and U. holubti Syd. which were described in 1895, 1898, 1911 and
1935 respectively have a number of points in common with the smuts
described in earlier years, though the last mentioned entirely destroys the
panicle while the others form sori in only some of the spikelets in an
inflorescence. Fullerton (1966) in a short report of his studies of soral
development of a smut in parts of the inflorescence of Hchinochloa colonum
referred that smut to Ustilago sphaerogena Burrill, using the work of
Fischer (1953) as a guide to its identity. Fullerton found that galls similar
to those in the floral parts were sometimes developed on the vegetative
parts of plants that had been inoculated with the ovary smut known to
him as U. sphaerogena. He also pointed out that the differences between
U. sphaerogena and U. crus-galli were differences of degree rather than of
kind and foreshadowed a taxonomic investigation of smuts of Hchinochloa.
Species of Ustilago on Echinochloa have been studied and the results
are reported in this paper. An account is given of the development and
structure of sori and of sporogenesis in both reproductive and vegetative
parts of the host. Using these and other data, a taxonomic revision of
certain smuts has been made.
PROCEEDINGS OF THE LINNEAN Society of NEw SourH WALES, VoL. 93, Part 3
282 A STUDY OF SOME SMUTS OF ECHINOCHLOA SPP.
It will be convenient in the following discussions of the smuts of
Echinochloa to refer to some of the specimens in terms related to the type
of sorus, ie., the symptoms, induced in the host and to the form of the
spines developed on the spore wall. Sori developed separately from various
organs of the floret are referred to as the U. sphaerogena form and sori
on vegetative parts or developed in the inflorescence as galls not specific to
particular organs are termed the U. crus-galli form. Spines on the spores are
referred to as being of the U. sphaerogena form or the U. crus-galli form when
they resemble the spines occurring on the spores in the type material of these
species.
MATERIALS AND MerrHops
Specimens of EHchinochloa colonum infected by a smut of the
Ustilago sphaerogena form were collected in the Rosewood district of
south-eastern Queensland during the late summer and autumn months (Feb—
May) of 1964 and 1965. In May, 1966, specimens were collected from the
Meandarra and Goondiwindi districts of Queensland. Some of the material
from these collections was dried, the remainder being preserved in Formalin
Acetic Alcohol (FAA).
In October, 1964 and September, 1965, seeds of Hchinochloa colonum
were dusted with spores of the U. sphaerogena form of smut and planted
in a garden plot at St. Lucia. A number of these plants developed sori.
A detailed study was made of the morphological and anatomical features
of the various soral types represented. Sori in different stages of development
were fixed in FAA. Material to be sectioned was dehydrated in a tertiary
butyl alcohol series and embedded in paraffin wax of MP 54-56° C. Blocks
were serially sectioned at 10u, sori being cut both longitudinally and
transversely. Two methods of staining were used, Safranin-Fast Green
(Johansen, 1940) and Periodic acid-Schiff’s Reagent (Conn, 1960) with a
Fast Green counterstain. Both methods were satisfactory for differentiating
mycelium and host tissues.
Details of spore formation were elucidated by examining the appropriate
parts of the serial sections by phase contrast microscopy. Some useful
supplementary information was obtained by hand teasing young sori, treating
the sporogenous hyphae with concentrated potassium hydroxide to remove
gelatinous material, mounting in clear lactophenol and examining by phase
contrast microscopy.
For the taxonomic work on the smuts of Echinochloa spp. specimens inow
a number of herbaria were obtained. These were supplemented by material
collected from plants in the field and smut-inoculated plants grown in
garden plots in the grounds of the University of Queensland at St. Lucia,
Brisbane. Spores for microscopic examination were mounted in Shear’s
mounting fluid (Graham, 1959). Herbaria from which specimens have been
made available are as follows: United States National Fungus Collection,
U.S.D.A., Beltsville (BPI) ; National Herbarium, Department of Agricultural
Technical Services, Republic of South Africa (PRE), New South Wales
Department of Agriculture, Rydalmere (DAR); Instituto de Botanica C.
Spegazzini, La Plata, Argentina (LPS), Commonwealth Mycological Institute
(IMT); Department of Botany, University of Queensland (BRIU).
MorpPHouocy or Sort
The sori on Hchinochloa spp. are readily detected because parts of the
plants where the fungus sporulates increase in size and bear numerous
hairs on the surface. The hirsute nature of the soral covering was noted
by Kunze (1830) when he described Ustilago tricophora, and similar
observations have been made Burrill (1888), Tracy and Earle (1895) and
R. A. FULLERTON AND R. F, N. LANGDON 283
Mundkur (1948) in their descriptions of various species of Ustilago which
occur on Hchinochloa spp. Magnus (1896) in describing the smut galls
on EH. crus-galli referred to the presence of a sheath of fungal hyphae
underlying the host tissue that covered the swollen areas. McAlpine (1910)
also has commented briefly on the structure of the galls studied by Magnus
but added no new information of any significance. The development of
sori in vestigial ovaries in staminate spikelets of Buchloe dactyloides infected
by Tilletia buchloeana was recorded by Norton (1896). Hansing and
Lefebvre (1941) described sori of Sphacelotheca sorgm and S. cruenta where
stamens as well as the Ovaries had been transformed by the smut and they
referred also to sori developing in the rudimentary ovaries of staminate
spikelets of Andropogon furcatus infected by Sorosporium everhartii. Fullerton
(1966) has shown that smuts of Echinochloa spp. which could be referred
to Ustilago sphaerogena were not confined to the ovary but spored in various
organs of the floret and in stems and leaves.
STRUCTURE OF SORI IN INFLORESCENCES
Since the form of the sori is influenced by the structure of the organs
or parts in which sporulation occurs, the morphology of floral parts of
Echinochloa colonum is described briefly here. Spikelets consist of two
florets, the lower floret being sterile and the upper one fertile. The sterile
floret has a lemma and a membranous palea only. The fertile floret has a
lemma which is hard and shiny with inrolled margins. With a thin palea
it encloses the floral organs. Small unicellular hairs occur on the outer
surfaces of the glumes, the sterile lemma and the pedicel of the spikelet
below the glumes. On the axes of the panicles are long, pointed, unicellular
hairs which arise in the epidermis.
The ovary is smooth, sub-globose and glistening white, about 0-5 mm.
long, with two styles and feathery stigmas. The ovary wall surrounding
the ovule is two cells thick. The ovule is anatropous with two integuments
surrounding the nucellus. There are three stamens and two _ lodicules.
The latter are flattened and fan-shaped, up to 0-5 mm. long, expanding at
the time of anthesis to 1-5 mm. diameter and becoming cup-shaped.
Macroscopic features of sori
Sori in inflorescences may be restricted to the floral parts of a minority
of spikelets, a condition which can be termed the Ustilago sphaerogena form
of smut expression (Plate xx1Ir).
Where sori replace organs of the floret, a spikelet bears a number of
globose, green, hispid sori up to 3 mm. diameter. In any infected spiklet
there are up to six sori of varying dimensions. When the maximum number
of sori is developed one sorus occupies the position of the ovary, three
are present in positions where stamens usually are placed, and a further
two occur lower down on the axis in the position normally occupied by
lodicules (Plate xx111). There is, however, great variation in this pattern. Not
uncommonly fusion of smutted floral organs leads to the formation of a
large and somewhat irregular sorus on the apical part of which may be
seen remnants of tissue recognizable as styles or stigmas, stamens or lodicules.
Sometimes parts of the lemma and palea are incorporated in such a sorus.
The sterile floret sometimes is changed, the palea being abnormally hairy
at the base. A small hispid structure, the origin of which could not be
determined, sometimes develops between the lemma and palea. All sori
retain their green colour until maturity, when the soral covering splits
irregularly to disclose a dark spore mass in which one or more columellae
become visible as the spores are dispersed.
284 A STUDY OF SOME SMUTS OF ECHINOCHLOA SPP.
A second sort of smut expression in inflorescences is that which might
be termed the Ustilago crus-galli form since sori occur in stems and leaves
and extend to inflorescences in some plants only. In the latter case almost
_ every spikelet in the inflorescence may be smutted and those not affected
are usually infertile. Various components of the spikelets may be affected
individually or in combinations to form a large number of irregularly shaped
sori. Sporulation in the sterile florets is common, lemmas and paleas being
sites for the formation of sori. Phyllody is a characteristic feature. This
is partly due to the extension of the distal ends of many sori into flattened
green leaf-like structures, but short vegetative shoots also grow from the
axis of the inflorescence. Unlike the sori of the U. sphaerogena form, these
sori do not maintain a regular shape for any one organ of origin and
there is considerable disparity between sori. Because of phyllody and the
range of plant structures involved it is difficult to identify with certainty
the origin of many of the individual sori (Plate xxiv).
Microscopic appearance of sori
The soral covering with layers of host and of fungal origin is a feature
common to all sori. The structure of the sori and of the collumellae contained
therein varies according to the part of the plant in which sporulation has
occurred.
The columella
The term columella is used in reference to the region where the fungus
and host tissues are in close contact. The columella in sori developed from
organs of the floret is a columnar structure extending upwards from the
base of the sorus and ending bluntly in the soral cavity. In sori in vegetative
tissues and sometimes in sori in inflorescences (the U. crus-galli form) the
columella is a pad of tissue with an undulating surface with which hyphae
are in close contact. In each case the columella is composed of cells of the
host (parenchyma and tracheal elements) and abundant intercellular and
intracellular hyphae.
The soral covering
The soral covering consists of two distinct layers, the outer one being
tissue of the host plant and the inner layer a fungal sheath (Plate xxiv). The
fungal sheath lining the soral cavity is a dense mass of interwoven hyphae
which can be separated from the host tissue by gentle manipulation with
needles. This sheath is present at an early stage of sporulation and remains
virtually unchanged in thickness as the sorus expands and matures. The
TaBLE 1
Thickness of layers in the soral covering
Thickness of Thickness
Type of sorus host tissue of fungal
layer layer
(All figures in microns)
Fully developed ovary sorus Bs 30-50 30-50
Dehiscing sorus in ovary ye BN 10-20 30-40
In stamen ape a Ms .. 50 (100 at cushions) 10-20
Fused ovary and stamen is a8 10-40 10-20
In lodicule at ra a3 ss 30-80 10--30
thickness of the host layers and the fungal layers in soral coverings varies,
as shown in Table 1. The host tissue has three to five layers of cells which
vary in size, ranging in diameter from 10 to 30u. In some places large
epidermal cells are grouped together to form cushions. It is from cells
MQ
fed O#
R. A. FULLERTON AND R. F. N. LANGDON
of these cushions that most of the hairs of the soral covering arise, although
soral hairs oecasionally develop from smaller epidermal cells. The hairs are
unicellular, elongate, tapering and sharply pointed structures. They vary
in length and usually reach 1 to 2 mm. with a basal diameter of 16 to 20p.
These hairs resemble those found on axes of the inflorescence and are much
larger than the hairs found on parts of normal spikelets. While spore
formation progresses the soral coverings increase to accommodate the
expanding spore mass. As the sorus approaches maturity, cells of the
host tissue distort and flatten as they dry and in some areas they may
peel off, exposing the fungal sheath which then is the only covering over
the spores.
Sori in ovaries
Sori developed from the ovary alone have in most cases a distinct
papilla situated on the distal end of the sorus and bearing two styles
and stigmas. A light coloured strip extending from this papilla down the
anterior side of the sorus can be seen in practically all sori of this type. In
this region there are two walls enclosing a flattened, elongate cavity.
The outer wall enclosing this cavity consists of three to four layers of
parenchymatous cells bounded inside and out by a distinct epidermis. There
is profuse hair development on both sides of the outer wall and also on the
epidermis of the inner wall.
Dissection showed that remnants of an ovule were often present in
this cavity which therefore has been interpreted as being the distorted
locule of the ovary. The form and the position of the ovule varied. In
some sori it was located in the papillate distal part of a sorus, somewhat
removed from the region of sporulation. Its structure then was approximately
that of the ovule in a non-infected ovary except that the integuments were
densely clothed on both sides with short, hyaline, unicellular hairs. In
other sori the ovule appeared to be placed directly on the wall of the sorus
and the normal morphological characters of the ovule were no longer evident
(Plate xxv, a, b).
In the early stages of development of sori in ovaries, mycelium sometimes
proliferates in the region of ovule attachment. In other cases the mycelium
grows into the funicle and may reach the base of the nucellus. The
consequent amount and type of distortion appear to be related to the position
in the ovary at which massing of mycelium occurs.
Sori in stamens
These sori were up to 2 mm. diameter, globose to fiask-shaped, often
surmounted by a filament up to 0-5 mm. long and bearing an anther. Such
anthers did contain pollen, although it was not determined whether this
pollen was fertile.
The first evidence of infection in filaments was a slight swelling and
marked hairiness close to the base of the filament. At a later stage a mass
of mycelium was found to be present inside the swollen portion. Further
growth resulted in the formation of a globose sorus with a typical cylindrical
columella inside (Plate xxvr).
Alteration of the staminal filament may extent for some distance above
the sorus proper, the remaining length of filament often being distinctly
swollen, tapering from sorus to anther and exhibiting marked hairiness.
Sori in lodicules
Sori in lodicules are up to 2 mm. diameter, but even in a single floret
there may be one much bigger than the other. The fungal sheath and its
286 A STUDY OF SOME SMUTS OF ECHINOCHLOA SPP.
overlying hispid covering are similar to those of sori in other organs. The
columella is broadly triangular in section and flattened in the plane of
the broad axis of the normal lodicule.
Other sori in wflorescences
The irregular sori of the Ustilago sphaerogena form originate in the
floral axis, and the regions of sporulation may include basal parts of
floral organs, lemmas and paleas. The soral cavity is continuous although
invaginations of the soral wall indicate where various host parts have grown
together. The columella of these compound sori is broad at the base and
divided apically, the points corresponding to the several components of
the sorus.
Some sori in inflorescences are not confined to the spikelets and resemble
the sori in vegetative parts of the plant. These sori of the Ustilago crus-galli
form, whether in inflorescences or in vegetative parts, have the soral covering
and the columella of the form already described. Other matters of structure
are discussed in a later section.
STRUCTURE OF SORI IN VEGETATIVE PARTS
Sori form as irregular blister-like swellings on various vegetative parts
of the plant (Plate vu a, 6). They are usually largest and most numerous,
often coalescing, in the nodal regions but occur commonly on the leaf bases,
leaf blades and internodes of the stem. The sori, which have green, hispid
coverings, vary from smoothly rounded to cerebriform. At nodes and on
leaf bases sori may reach a size of 1:0—2:0 cm. diameter. On leaf blades
and in internodal regions sori are usually much smaller, 1-several mm.
diameter, occurring as isolated blisters or by coalescence forming sori which
lie along the leaf or stem. Normal inflorescences are commonly produced
by the infected culms. The presence of smut in nodal regions of these
culms may stimulate development of axillary buds leading to a rosette
of short, leafy axillary shoots emanating from a complex mass of sori.
Inflorescences are seldom produced from such shoots. The microscopic
structure of the sori is the same as that of galls in inflorescences except
for certain details of the form of the columella. The points of difference
are mentioned below.
SPORE FORMATION
Kukkonen and Vaissalo (1964) have reviewed the rather limited
information that there is on spore formation in smut fungi. Their own
electron microscope studies of sporogenesis in Anthracoidea aspera confirm
earlier work on species of Ustilago by De Bary (1887), Lutman (1910) and
Hutchins and Lutman (1938) who found that spore initials were embedded
in an almost homogeneous gelatinous mass produced by swelling of the
sporogenous hyphae. Magnus (1896) and McAlpine (1910) have recorded
some details of development of Ustilago crus-galli. In that smut gelatinization
of hyphae was observed, and weight was given to the supposed basipetal
formation of spores in the sporogenous hyphae. The latter observation was
the reason for these authors’ inclusion of the species in the genus Cintractia.
De Bary (1887) defined the genus Sphacelotheca, emphasising the columella
and the fungal sheath surrounding the spore mass as characters by which
it was distinguished from Ustilago. He said that the development and
mature structure of the spore mass of Sphacelotheca were the same as
those of Ustilago. For the latter he described the gelatinization of hyphae
synchronously with or even before the basipetal development of the spores.
R. A. FULLERTON AND R. FF. N. LANGDON 287
During the present investigation the modes of spore formation of sori
of the Ustilago sphaerogena and U. crus-galli forms respectively were
found to be essentially the same. Sori of the former tend to be symmetrical
about the columella while the latter usually have the columella oriented
so that the sporulation is above it and not around it. There is a continual
production of hyphae at the columella. The walls of these hyphae become
gelatinized and hyphal characters disappear. Spore initials then appear in
the gelatinous matrix a short distance from the columella, and when first
visible are opaque bodies about 3 in diameter, almost indistinguishable
from the rest of the matrix. The spore initials then rapidly expand to a
size approaching that of mature spores. Concurrent with the development
and expansion of spore initials is the formation of dense echinulations which
outline the spores, no spore walls being visible at this stage. Soon after
the appearance of echinulations the spore walls become well defined. At
first hyaline, they soon become pigmented. As the initials are expanding
and maturing, the gelatinous matrix gradually disappears until the spores
are connected by thin strands of gelatinous material. In young sori spore
masses are agglutinated and possibly the cementing material may be remnants
of this matrix. In older sori spores become powdery.
The production of sporogenous mycelium in young sori proceeds at a
greater rate than its gelatinization. Thus the columella becomes surrounded
by mycelium which is hyaline, thick walled, septate, much branched and
1-0-3-0u in diameter. It closely resembles the mycelium of the sheath that
surrounds the spores.
The gelatinization of sporogenous mycelium that grows around or
above the columella tends to be concentrated in a number of discrete regions.
The continued growth of mycelium between the regions of rapid gelatinization
produces hyphal columns which extend into the developing sorus, the columns
being more numerous in the sori in vegetative parts than in floral sori.
These columns in which the hyphae lie parallel to one another are associated
with crests on the undulating surface of the columella (Plate xxvir). Gelatini-
zation of hyphae of the columns later reduces considerably the extent to
which the columns penetrate out into the mature spore mass.
During development of the sorus, spores in all stages of formation
can be found. When production of sporogenous mycelium ceases, spore
formation continues and mature spores are finally developed very close to
the columella.
TAXONOMY
Ustilago tricophora (Link) Kunze. Flora, 13, 869. 1830. Caecoma
tricophorum Link in Willdenow, Sp. Pl., 6 (2), 3. 1825. Ustilago sphaerogena
Burrill in Saccardo, Sylloge Fung., 7, 468. 1888. Ustilago crus-galli Tracy
& Earle, Bull. Torrey bot. Club, 22, 175. 1895. Ustilago panici-frumentacer
Brefeld, Unters. Gersammt. Mykol., 12, 103. 1895. Cintractia seymouriana
Magnus, Ber. dt. bot. Ges., 14, 217. 1896. Ointractia crus-galli (Tracy
& Earle) Magnus, Ber. dt. bot. Ges., 14, 392. 1896. Ustilago globigena Speg.,
Anal. Mus. nac., 6, 208. 1898. Cintractia sphaerogena (Burrill) Hume, Proc.
Towa Acad. Sci., 9, 233, 1902. Ustilago tricophora (Link) Kunze var. pacifica
Lavroff, Trudy biol. nauchno-issled. Inst. tomsk. gos. Univ., 2, 9. 1936.
Ustilago tricophora (Link) Kunze var. crus-galli (Tracy & Earle) Lavroff,
Trudy biol. nauchno-issled. Inst. tomsk. gos. Univ., 2, 9. 1936.
Sori in organs of the floret, in parts of the inflorescence or in vegetative
parts, variable in size and form, with a hispid covering of host tissue. Spore
mass agglutinated at first, later pulverulent, surrounded by a fungal sheath
288 A STUDY OF SOME SMUTS OF ECHINOCHLOA SPP.
up to 50u thick. Spores globose, subglobose or ellipsoid, dark, 6-14, at
greatest diameter, ornamented with spines varying in density of distribution,
length and shape.
Specimens examined: On Echinochloa colonum (L.) Link, Egypt,
- Ehrenberg, no date (IMI); Sudan, 8S. A. J. Tarr, 12. x. 1954 (IMI 59760) ;
Louisiana, U.S.A., Atkinson & Forbes, 28. viii. 1936 (BPI); Louisiana,
U.S.A., C. R. Ball, 16. viii. 1898 (BPI); Louisiana, U.S.A., I. L. Forbes,
11635 Ibs "1940 (BPI); Cuba, F. S. Earle, 9. x. 1924 (BPI) ; Amberley, Qld.,
R. F. N. Langdon, 29. iii. 1947 (BRIU 415) ; Lawes, Qld., W. J. Bissett, 28.
li. 1941 (BRIU 501) ; Nambour, Qld., J. C. Johnson, 26. 111. 1951 (BRIU 270) :
Cleveland, Qld., O. R. Byrne, 1951 (BRIU 1209); St. Lucia, Qld. R. A.
Fullerton, iii. 1966 eR 2283), ii. 1966 (BRIU 2284), iii-iv. 1966 (BRIU
2286), 3. iv. 1966 (BRIU 2287); Meandarra, Qld., R. A. Fullerton, 1. v. 1966
(BRIU 2288) ; Goondiwindi, Qld., R. A. Fullerton, 2. v. 1966 (BRIU 2289,
2290) ; Pilliga, N.S.W., J. A. O’Reilly, vi. 1958 (DAR 4863). On Echinochloa
turneriana Domin., Windorah, Qld., P. J. Skerman, 26. vii. 1946 (BRIU 399),
24. vi. 1949 (BRIU 529). On Echinochloa holubii (Stapf) Stapf, Transvaal,
Sth, Africa, I. B. Davy, 15. 11 1912 (PRE 2247, BPI) (TYEE, Ustilago
crus-galli Tracy & Earle var. minor Zundel). On Echinoch loa walteri (Purch)
Heller, Connecticut, U.S.A., G. M. Reid, 14. ix. 1919 (BPI). On Hehinochloa
crus-galli (L.) Beauv., Utah, U.S.A., Tracy & Evans, no. 651, 8. x. 1887
(BPI) (TYPE, Ustilago crus-galli Tracy & Earle); Dlinois, U.S.A., A. B.
Seymour, no. 1892, ix. 1884 (BPI) (TYPE, Ustilago sphaerogena Burrill) ;
Argentina, ©. Spegazzini, no. 3025, 13. v. 1917 (LPS) (Ustilago globigena
Speg., det. C. Spegazzini); Nanking, China, F. L. Tai, 3. x. 1929 (BPI);
Japan, quarantine interception Be Beatle, Wk S.A., ili. 1941 (BPI); Mandalay,
Burma, A. McKerrel, 17. viii. 2 (BPI); Arizona, U.S.A., D. F. Cook,
22 ep ixe 1923 (BRM) Gieene Te W. W. Mackie, xii. 1928 (BPI):
Colorado, U.S.A., A. S. Hitchcock, 28. viii. 1906 (BPI); Colorado, U.S.A.,
K. Bartholomew, 4. ix. 1914 (BPI); Connecticut, U.S.A., G. P. Clinton, 22.
ixsal90G) (GEE) aConnectieutn WU SiAsy Gai: Zundel, 30. ix. 1926 (BPI);
Florida, U.S.A., H. W. Wedgeworth, 3. vi. 1940 (BPI) ; Illinois, U.S.A.,
M. B. Waite, 2. x. 1888 (BPI); Indiana, U.S.A., F. D. Fromme, 1. x.
19138 (BPI); Iowa, U.S.A., L. H. Pammel, 20. ix. 1909 (BPI); Maryland,
WES ZAC Hy Ee McKinney, 25. x. 1944 (BPI); Massachusetts, U.S.A., A. B.
Seymour, 10. ix. 1910 (BPI); Minnesota, U.S.A., D. Griffiths, viii. 1896
(BPI); Missouri, U.S.A., J. B. Norton, ix. 1896 (BPI); Nebraska, U.S.A.,
TS SAG Williams, 11. ix. 1890 (BPI); Nevada, U.S.A., O. F. Smith, viii. 1940
(BPI); New Jersey, U.S.A., F. L. Scribner, 24. ix. 1880 (BPI); New Mexico,
U.S.A., E. W. D. Holway, 18. ix. 1896 (BPI); New York, U.S.A., R. Latham,
x 1916 (BRD). Oklahoma, WeS.A) (W.,He. laong, 28) oxi 909 Gaia
Oregon, U.S.A:, J. R. Kienholz, 15. ix. 1937 (BPI); Pennsylvania, U.S.A.,
G. L.-Zundel, 22. ix. 1988 (BPI); Utah, U.S.A., A. D. Garrett, 16. x. 1904
(BPI); Virginia, U.S.A., P. Klaphaak, 12. x. 1922 (BPI); Washington,
U.S.A., W. N. Suksdorf, 3. x. 1894 (BPI); Washington, D.C., U.S.A., M. B.
Waite, 30. x. 1888 (BPI); Wyoming, U.S.A., A. Nelson, 27. viii. 1904 (BPI) ;
Spain, collector not stated, viii. 1946 (BPI); Morocco, G. Malencon, 9. xi.
1932 (BPI); Nigeria, A. Thompson, 2. iv. 1920 (BPI); Cape Province, South
Africa, A. O. D. Mogg, 7. iii. 1934 (PRE 27384: and BPTI, as Ustilago erus-galli
var. minor Zundel, det. Zundel) ; Argentine, T. Rojas, v. 1906 (BPI); New
York, U.S.A., R. Latham, x. 1922, and D. Reddick, 25. x. 1941 (BPE as
Ustilago crus-galli var. minor Zundel, det. Zundel) ; Transvaal, South Africa,
A. O. D. Mogg, 3. vii. 1934 (BPI, as Ustilago crus-galli var. minor Zundel,
det. Zundel); Richmond, N.S.W., W. M. Carne, iii. 1911 (DAR 721), A.
Murphy, v. 1961 (DAR 6152); Yenda, N.S.W., collector not stated, iv. 1934
R. A. FULLERTON AND R. F. N. LANGDON 289
(DAR 1002); Yanco, N.S..W., P. Kable, iii. 1962 (DAR 6981); Baulkham
Hills, N.S.W., collector not stated, 1956 (DAR 4862). On Hchinochloa crus-
gallt (L.) Beauy. var. frumentaceae W. F. Wight, Aspley, Qld., J. G. Morris,
iv. 1952 (BRIU 583).
In addition to the specimens already listed, 32 other specimens on
Echinochloa crus-galli from localities in 14 States of the United States of
America were examined. These fungi did not differ significantly from the
collections on that grass from the U.S.A. which have been formally listed
above.
Notes on specimens examined
(a) Type specimens
Ustilago sphaerogena Burrill. The specimen consists of two smutted
spikelets. Floral parts are affected individually. In one spikelet there is a
large sorus in the position of the ovary, bearing remnants of the styles on
its papillate apex. Three smaller sori surround the large central sorus,
occupying the positions of the stamens. Each of these three sori bears an
anther on a thin projection from the apex. Two small sori occupy the
positions of the lodicules. All sori are covered by hairy tissue of the host.
The other spikelet bears several sori occupying the positions of the floral
parts. The spores are globose to subglobose, 6-124 diameter, densely and
sharply echinulate (Plate xxvii1).
Ustilago crus-galli Tracy & Earle. The specimen is a small length
of stem with a single node and part of a leaf. An axillary shoot is
developed from the node. Sori are developed in two hairy leaf galls each
about 2-0 mm. diameter. The spores are globose to subglobose, 9-0-13-5p
diameter, echinulate. The spines are broader at the base (i.e. blunter) and
less densely crowded than in U. sphaerogena (Plate xxvitt).
Ustilago crus-galli Tracy & Earle var. minor Zundel. The spores of
this specimen are not distinguishable from those of type material of
U. crus-galli.
(b) Authentic specimens
' Ustilago globigena Speg. The specimen is part of an inflorescence with
a few smutted spikelets. The sori are developed in floral parts and resemble
closely the sori found in the type specimen of U. sphaerogena. The spores
are not smooth, as stated by Spegazzini (1899), but are echinulate and
as suggested by Hirschhorn (1939) are very similar to those of U. sphaerogena.
(c) Other specimens
Ustilago tricophora (link) Kunze. Type material could not be located.
By courtesy of the Director of the Commonwealth Mycological Institute we
have been able to view a slide made by one of his staff from a smutted
specimen of Hchinochloa colonum which is now in a phanerogamic herbarium
(not specified). The specimen had been collected in Egypt by Ehrenberg,
who was also the collector of the material from Egypt on which Ustilago
tricophora is based. There is a strong possibility that Ehrenberg’s -collection
from Egypt went partly to Link and partly to a phanerogamic herbarium,
especially as the inflorescence smut of Hchinochloa affects only a small
proportion of the spikelets. The spores of this specimen are globose
to subglobose, 9-11» diameter and densely echinulate. The spines are slightly
coarser and not quite as long as the spines of Ustilago sphaerogena. A
specimen of smut on Hchinochloa colonum from the Sudan is very similar
to type material of Ustilago sphaerogena, as also is smut on Echinochloa
erus-galli from Cape Province, Union of South Africa. It is believed that
290 A STUDY OF SOME SMUTS OF ECHINOCHLOA SPP.
these specimens from Africa, including one in the type host from the type
locality of Ustilago tricophora, can be accepted as reliable evidence as to
the characteristics of Ustilago tricophora.
Ustilago sphaerogena Burrill and U. crus-galli Tracy & Earle. A
majority of the specimens received under the name Ustilago sphaerogena
bore sori in the floral parts which were the same as those seen in the type
specimen of this fungus. The spores also were characteristic of that species.
Similarly, most of the specimens which were labelled Ustilago crus-galli had
the soral and spore characters of the type material of that species. The
dimensions of the spores in some specimens varied a little from those of the
type specimens of these two species. Specimens of the U. sphaerogena form
in all except three cases had spores which were within the limits of size
of spores of the type specimen, and these exceptions exceeded the upper
limit by only one micron. With U. crus-galli material the type specimen
had spores ranging from 9 to 13-54 diameter but in a majority of specimens
spores with lower limits down to 6 were found. A few specimens included
spores which had their longest dimensions one or two microns greater than
what were observed in the type specimen. Spores which exhibited these
deviations from the limits of size of the type specimen were nearly all
markedly subglobose or tending to ellipsoid. For both species the variations
in dimensions are not regarded as being of any significance. All specimens
include spores with dimensions within the range exhibited by the type
specimens.
The type specimen of Ustilago sphaerogena exemplifies a form of
Echinochloa smut where sori are developed in the floral parts alone. The
ornamentation of the spores of certain specimens with these symptoms was
not typical of Ustilago sphaerogena. In some the spines were less densely
spaced and the form of the spines tended towards that found in spores of
U. crus-galli, i.e., somewhat shorter and more broadly based than in U.
sphaerogena. Three specimens from Louisiana, U.S.A. (collected by Ball
in 1898, Atkinson and Forbes in 1936 and Forbes in 1940 respectively)
exhibited this intermediate condition of the spines. In specimens from Cuba
(coll. Earle, 1924), China (coll. Tai, 1929) and Spain (collector not stated,
1946), the ornamentation of the spores was typical of that found in spores
of type material of U. crus-galli. A specimen from Nigeria (coll. Thompson,
1920) had spores with the U. crus-galli form of spines but with the spines
more sparsely placed than in U. crus-galli. A few spores of this specimen
were smooth. A collection from Oregon, U.S.A., by Kienholz in 19387 which
had the U. sphaerogena form of sorus had very short, broadly based and
sparsely placed spines which could be considered to be within the range
of variation of U. crus-galli, and indeed this specimen, identified by R.
Sprague, came to us under the name U. crus-galli.
Two specimens with stem sori typical of Ustilago crus-galli (Burma,
coll. MeKerrel, 1922, and Connecticut, U.S.A., coll. Clinton, 1906) had spores
with ornamentation tending towards the form typical of U. sphaerogena.
The former was labelled U. sphaerogena, the latter U. crus-galli. Both are
intermediate in form between the “typical” conditions of those species.
Specimens of smut on Hehinochloa colonum from Queensland included
some specimens collected in the countryside and others from plants grown
in plots at St. Lucia. Most plants from the field bore sori of the Ustilago
sphaerogena form only and had spores which conformed to those described
for U. sphaerogena. Exceptions to this were BRIU 2288 which had sori
on vegetative parts only, BRIU 2289 which had sori in both vegetative parts
and inflorescences, the sori in the latter being of two kinds, the U. crus-galli
form and the U. sphaerogena form, and BRIU 2290 with sori of the U.
R. A. FULLERTON AND R. F. N. LANGDON 291
crus-galli form in both vegetative parts and inflorescences. Plants grown
from smut-inoculated seed also included some which bore sori of both the
U. crus-galli and the U. sphaerogena forms (BRIU 2283, 2284, 2286) and one
with the U. crus-galli form of sorus in vegetative tissues only (BRIU 2287).
The ornamentation of the spores in these specimens was variable. In
some sori forms of spine were found which were intermediate between the
U. sphaerogena and U. crus-galli forms in width and in distribution on the
spore surface. In others, spores of the typical U. crus-galli form were found.
In essence, the spores and the sori on plants inoculated with spores from
the U. sphaerogena form of sorus showed variability covering the range
of characters which earlier workers had used to delimit two species, namely
U. sphaerogena and U. crus-galli.
Notes on synonymy
Three names have been listed as synonyms of Ustilago tricophora, namely
U. panici-frumentacei, U. tricophora var. pacifica and U. tricophora var.
crus-galli, although type or authentic specimens have not been seen. Mundkur
(1943) has studied U. panici-frumentacei and considered its relationship to
U. tricophora. A small difference in spore size was the only character by
which Mundkur could differentiate these two species. Our present studies
do not support a concept of species differentiation by size differences of
about one micron. Lavroff (1936) included U. panici-frumentacei with
U. tricophora, giving it varietal status in the latter species. He also reduced
U. crus-galli to the status of a variety of U. tricophora (Petrak, 1950).
Mundkur (1943) after examining type material of U. paradoxa reported
that its spores were entirely smooth. He noted that its mode of germination
differs from that of U. crus-galli and U. panici-frumentacei. In our studies
only two specimens, one from Nigeria (BPI) and the other from Queensland
(BRIU 2284), have exhibited spores lacking spines. In these fungi the
smooth spores were exceptional, the great majority being echinulate. U.
paradoxa is accepted as being distinct from U. tricophora although its habit
of. attacking only some spikelets in an inflorescence and its hispid soral
covering indicate its possible relationship to U. tricophora.
DISCUSSION
Several species of Ustilago have been described from Hchinochloa from
various parts of the world. One of them, U. holubii, destroys the inflorescence
and the sorus may extend into the stem below. This characteristic and
the smooth to minutely verruculose spores distinguish it from other smuts
of these grasses. Common to the rest of the species of Ustilago described
from Hchinochloa is a hispid membrane, developed from tissues of the host,
covering the fungal structures in the region of sporulation. In a taxonomic
study of smuts of this kind we have retained Ustilago paradogxa, which has
smooth spores, and have grouped the other smuts to make a single species,
Ustilago tricophora. The description of the latter species has been amended
in this paper and it now includes all those smuts of Hchinochloa which
form sori with coverings formed conjointly of fungal tissue and hispid tissue
of host origin and which have spores ornamented by spines.
Certain smuts of Echinochloa may evoke similar host reactions which
are reflected by the uniformity of the host tissues that cover sori and by
resemblances between sori on various parts of the plant. Apparent differences
in soral structure, e.g. in shape and size, are related to the organ or part
292 A STUDY OF SOME SMUTS OF ECHINOCHLOA SPP.
of the plant in which sporulation has occurred. Similarities in soral characters
are not in themselves indicative of a taxonomic relationship between smuts.
Evidence for such a relationship for the smuts now referred to Ustilago
tricophora is to be found in the experimental work on inoculation of plants
with smut, in patterns of sporogenesis in the different forms of sori, and
in comparisons of spore characters of a large number of herbarium specimens.
It has been shown that when Echinochloa colonum was inoculated with
spores from the Ustilago sphaerogena form of sorus individual plants bearing
sori of both the U. sphaerogena and the U. crus-galli forms could be found.
In these sori supposedly typical of two different species of smut, there were
marked similarities in the development of fungal structures, e.g., columella
and fungal sheath, and in sporogenesis. In the inoculated plants as well as
in the herbarium specimens, spores representing the intergrades between the
U. sphaerogena form and the U. crus-galli form were found. The extreme
cases were the occurrence of the U. crus-galli form of spores in sori in the
U. sphaerogena form of sorus, and vice versa.
Fullerton (1966), who reported on the structure of the sori of the
Ustilago sphaerogena form of smut of Echinochloa colonum suggested that
Sphacelotheca might be a genus more appropriate than Ustilago for the
species of smut he studied. The basis for that suggestion was that the
smut of Echinochloa appeared to have the characteristics of spores enclosed
by a fungal sheath and basipetal formation of spores around a columella
which in recent years have been accepted as characteristic of the genus
Sphacelotheca (Fischer, 1953; Fischer and Holton, 1957). Hirschhorn (1939)
and McAlpine (1910) have doubted the validity of Sphacelotheca as a genus
distinct from Ustilago. McAlpine noted that a columella occurs in other
smuts such as species of Cintractia and Sorosporiwm, and that there are
“. . various gradations in the formation of a fungus membrane enclosing
the spores ...”. He indicated that there were difficulties in deciding whether
a Sheath should be regarded as evanescent or persistent. Concerning the
other characters for distinguishing Sphacelotheca from Ustilago, namely the
basipetal formation of spores in the former which is lacking in the latter,
there seems to be no grounds for accepting such a distinction. De Bary
(1887) declared that in the smut on which Sphacelotheca was based, namely
Ustilago hydropiperis, “. . . the development and mature structure of the
Spore-mass are the same as those of Ustilago ...”. In the sections cut from
Ustilago tricophora in various stages of development we have noted that for
some time there is production of sporogenous hyphae around the columella.
Spores develop from the first-formed hyphae and are pushed outwards as
more sporogenous hyphae grow. Spore formation is a continuous process
that occurs progressively while sporogenous hyphae are forming rather
than being the basipetal formation of spores from a mass of pre-formed
hyphae, progressing from the region of the fungal sheath towards the
columella. The basipetal formation of spores often quoted as a characteristic
of the genus Sphacelotheca may be more apparent than real. Further studies
of sporogenesis in smut fungi which at present are included in Sphacelotheca
are in progress. For the present no change in the generic position of the
species tricophora will be made.
Acknowledgements
The authors acknowledge with thanks the financial assistance provided
by The University of Queensland Research Committee and the cooperation
of the directors of herbaria from which specimens were borrowed.
Proc. Linn. Soc. N.S.W., Vol. 93, Part 3 PLATE XXIII
|
ses
Top.—Echinochola ‘colonum with Ustilago sphaerogena form of smut.
Bottom.—Sori developed in floral parts of Hehinochloa colonun.
Proc. Linn. Soc. N.S.W., Vol. 93, Part 3
Top.—Echinochloa colonun with Ustilago crus-galli
form of smut.
Bottom.—Section showing features of soral covering
of sori on Hehinochloa colonun.
PLATE
XXIV
2 F re
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PLATE xxv
Proc. Linn. Soc. N.S.W., Vol. 93, Part 3
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Proc. Linn. Soc. N.S.W., Vol. 98, Part 3
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Longitudinal section of sorus in staminal filament
of Hchinochloa colonum.
PLATE XXVI
i
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PLATE XXVII
Soc. N.S.W., Vol. 93, Part 3
Proc. Linn.
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0109 DOLYIOWLYOA JO SyVeeYys Jes] pue sapou uo TloS—'aprs play
Proc. Linn.
Soc. N.S.W., Vol. 93, Part 3
PLATE XxXvII
les and leaf sheaths of Hchinochloa colonurm.
1 on nod
Section through sorus on stem of Hchinochloa colonum.
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PLATE XXVIII
Proc. Linn. Soc. N.S.W., Vol. 98, Part 3
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WIOJ DUAHoLaDyds O6D1178Q 94} Jo satodg—‘aprs yfaT
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R. A. FULLERTON AND R. F. N. LANGDON 293
References
ANON., 1958.—Plant disease survey for twelve months ending 30 June 1958. (New South
Wales Department of Agriculture, Sydney.)
BurRiLL, T. J., 1888.—In Saccardo, Sylloge Fung., 7: 468.
Conn, H. J., 1960.—‘‘Staining procedures used by the Biological Stain Commission’, 2nd
Ed. (Williams and Wilkins, Baltimore.)
DE Bary, A., 1887.—‘‘Comparative morphology and biology of the fungi, mycetozoa and
bacteria.” (Clarendon Press, Oxford.)
FISCHER, G. W., 1953.—‘“Manual of the North American Smut Fungi.” (Ronald Press,
New York.) j
: , and Hotton, C. 8., 1957—‘“‘Biology and control of the smut fungi.” (Ronald
Press, New York.) 5
FULLERTON, R. A., 1966.—The structure of sori of Ustilago sphaerogena Burrill. Aust.
J. SCi., 28: 435-436.
GRAHAM, S. O., 1959.—The effects of various reagents, mounting media, and dyes on the
teliospore walls of Tilletia contraversa Kuhn. Mycologia, 51: 477-491.
HANSING, EH. D., and LEFEBRE, C. L., 1941.—Smut sori from ovarial and staminal tissues
of certain grasses. Phytopathology, 31: 1043-1046.
HirscHuHorn, H., 1939.—lLas especies del genero ‘“Ustilago” en la Argentina. Darwiniana,
3: 347-418.
Hutcuins, H. L., and Lurman, B. F., 1938—Spine development on the spores of Ustilago
zeae. Phytopathology, 28:. 859-860.
JOHANSEN, D. A., 1940.—‘‘Plant microtechnique.” (McGraw-Hill, New York.)
KUKKONEN, I., and VAISSALO, T., 1964.—An electron microscope study of spore formation
ina smut. Ann. Bot. Fenn., 1: 236-249.
Kunze, G., 1830— Flora, 13: 369.
LAvRorr, N. N., 1936—New and less common smut fungi of the Ustilaginaceae in
Northern and Central Asia. Trudy biol. nauchno-issled. Inst. tomsk. gos. Univ.,
2: 9. (Cited in Rev. appl. Mycol., 16: 342, 1937.)
LUTMAN, B. F., 1910.—Some contributions to the life history and cytology of the smuts.
Trans. Wis. Acad. Sci. Arts Lett., 16: 1191-1244.
Hine nordamerikanische Ustilaginee auf Panicum crus-galli. Ber. dt.
bot. ’ Ges., 14: oes D2
McALPINE, D., 1910.—‘‘The Smuts of Australia. ” (Government Printer, Melbourne.)
Munpkur, B. B., 1943.—Studies in Indian cereal smuts. VI. The smuts on sawam
(Echinochloa frumentacea). Indian J. agric. Sci., 13: 631-633.
Norton, J. B. S., 1896.—A study of the Kansas Ustilagineae especially with regard to
their germination. Trans. Acad. Sci. St. Lowis, 7: 229-241.
PETRAK, F., 1950.— ‘Index of Fungi, 1936-1939.” (Commonwealth Mycological Institute,
Kew.)
SPEGAZZINI, C., 1898—Fungi argentini novi vel critici. An. Mus. nac., 6: 81-367.
Tracy, S. M., and Harte, F. S., 1895—New species of parasitic fungi. Bull. Torrey bot.
Clubs 222 1b:
HXPLANATION OF PLATES
Plate xxiIr
Top. Hchinochloa colonum with Ustilago sphaerogena form of smut expression in floral
parts (x 8).
Bottom. Sori developed in floral parts of Hchinochloa colonum (x 14) (a) in ovary;
(0) in staminal filaments, with anthers still attached; (c) in lodicules; (d@) upper
glume and fertile lemma.
Plate xxiv
Top. Echinochloa colonum with Ustilago crus-galli form of smut expression in
inflorescence (x 3).
Bottom. Section showing features of soral covering of sori on Hchinochloa colonum
(x 160). (a) epidermal hairs; (0) host tissue; (c) fungal sheath; (d) spore mass.
Plate xxv
Left side. Longitudinal section of sorus in ovary of Echinochloa colonum with ovule in
papilla above sorus (x 35).
Right side. Longitudinal section of sorus in ovary of Hchinochloa colonum with ovule
on wall of sorus (x 30).
Plate xxvI
Longitudinal section of sorus in staminal filament of Echinochloa colonum (x 40).
Plate xxvilI
Left side. Sori on nodes and leaf sheaths of Echinochloa colonum (x 8).
Right side. Section through sorus on stem of Echinochloa colonum (x 30).
Plate xxvIIr
Left side. Spores of the Ustilago sphaerogena form (x 1000).
Right side. Spores of the Ustilago crus-galli form (x 1200).
A VIVIPAROUS SPECIES OF PATIRIELLA (ASTEROIDEA,
ASTERINIDAE) FROM TASMANIA
A. J. DARTNALL
The Tasmanian Museum, Hobart, Tasmania
(Plate xxix)
[Read 25th September, 1968]
Synopsis
A viviparous species of asterinid sea star of the genus Patiriella is described.
Attention is drawn to its restricted distribution in the littoral of S.E. Tasmania and
the significance of its mode of development is noted. Some comparisons are drawn with
other Australian species of Patiriella.
INTRODUCTION
The new species of Patiriella discussed in this paper was known formerly
from specimens held by the Tasmanian Museum and the Zoology Department
of the University of Tasmania. The specimens had been attributed to
Patiriella eaigua (Lamarck), 1816 and Asterina scobinata Livingstone, 1933.
Class STELLEROIDA
Subclass ASTEROIDEA
Family AstTerinipan Gray, 1840
Genus Patiriella Verrill, 1913
PATIRIELLA VIVIPARA, Sp. NOV.
(Plate xx1x)
Description of holotype—A small asterinid sea star with five arms. The
interradial edge is hardly concave; the entire animal having the appearance
of a slightly rounded pentagon.
The abactinal, radial area is covered with imbricate, crescentic plates.
A papular pore is enclosed by the concavity of each crescentic plate. The
papulae are arranged in five (5) rows on each side of the abactinal axis
of the arms. Papulae are absent from most of the interradial area of the
abactinal surface and the plates of this area are without a papular notch.
The abactinal plates carry 4 to 13 cylindrical, slightly capitate, granular
spinelets. The number of spinelets on each abactinal plate increases towards
the centre of the disc.
The madreporite is sub-triangular, channelled and perforate.
The anus is protected by six (6) spinelets inclined obliquely over the
aperture.
The inferomarginal plates are distinct, each carrying four (4) spinelets
on the outer margin, and form a conspicuous edge to the actinal surface.
The actinal plates are imbricate and arranged in chevrons in the
interradial area. Each plate is granular and carries one (1) bluntly pointed
spine. Towards the disc margin two (2) spines are present on some plates.
The adambulacral plates bear two (2) furrow spines and one (1)
subambulacral spine. The adambulacral plate proximal to the oral plate
carries three (8) furrow spines. The furrow spines are basally webbed,
broad and bluntly pointed. Each oral plate carries six (6) oral spines (the
largest being the innermost) and one (1) suboral spine.
Colour in life-—Life colour is a constant orange-yellow, slightly lighter
on the actinal surface.
Type material.—Holotype. R=85 mm., r=6-8 mm., Midway Point,
Pittwater, Tasmania (42°49’ S., 147°31’ E.). Littoral rock shelf. 238. iv.
1967. Collected, A. J. Dartnall, Tasmanian Museum Reg. No., H871.
PROCEEDINGS OF THE LINNEAN Society or NEw SoutH WALES, Vow. 93, Part 3
A. J. DARTNALL I95
Paratype series —Thirty-three (33) specimens ranging in size from
R=1-:0 mm. to R=105 mm. Two individuals fixed with emerging young.
(All are spirit specimens.) Locality details as for holotype. Tasmanian
Museum Reg. No. H372.
Distribution and Habitat—Patiriella vivipara is known from four
localities extant — all in south-eastern Tasmania — and is probably extinct
in a fifth locality due to recent modification of the foreshore (see Fig. 1).
Figure 1. Distribution of P. vivipara, sp. nov., in S.E. Tasmania. Solid blocks show
sites where the species is known to be extant; the cross the station where it occurred
recently, but from where it is absent now.
The species is found on gently sloping, sheltered shores, characterized by
rocks, often no more than a foot high, littered on a rock platform which
gives way to sand in the lower littoral. It is found under rocks in the
Galeolaria zone of the shore and it can be demonstrated to be more negatively
phototatic than both Patiriella eaigua and Patiriella regularis (Verrill),
1867.
Remarks.—Interest in this species stems from its viviparous habit. The
young develop in sacs derived from the gonads and when a size of 1-2 mm.
R is attained they rupture the incubatory sac, enter the coelom and emerge
between the abactinal ‘plates of the adult. P. vivipara breeds throughout
the year.
Retention of young during embryonic development in specially adapted
structures within or upon the body of the parent is common amongst
echinoderms. Boolootian (1966) lists thirty-three species of Asteroidea which
exhibit brooding habits, coelomic and ovarian incubation is known among
the Holothuroidea and bursal incubation in the Ophiuroidea.
P. vivipara is the only asteroid known to exhibit embryonic development
in a sac derived from the gonad and adds another reproductive variant to
the family Asterinidae.
A ffinities—Patiriella vivipara is very similar to P. eaigua. Both species
possess six oral spines (sometimes 5 or 7) at a size of 10-11 mm. R and
differences of ambulacral, actinal and abactinal spinulation and plate
arrangement, if any, are not distinct. Juvenile examples of P. regularis,
which may be confused with both P. vivipara and P. exigua in Tasmanian
waters, can be separated by the smaller number of oral spines (5 at 10-11
mm. R). (See Table 1.)
P. vivipara attains maximum size at 14 mm. R, this being exceeded by
all the Australian species of Patiriella.
296 A VIVIPAROUS SPECIES OF PATIRIELLA (ASTEROIDEA, ASTERINIDAE)
Reproductive characteristics may be used to separate the species and
P. vivipara is distinguished from other members of the genus, particularly
P. exigua, primarily on this basis. Amongst the species available for study
TABLE |
Some characteristics of species of Patiriella in Australia
Number
Number Number of Spines
| Species of of Oral per Origin of data
. rays spines actinal
plate
P. vivipara 5 6* 1 Tasmanian Museum collections
P. exigua 5 6* 1 Bp » >
P. regularis 5 5* 1 ay a Je
P. nigra 5 T 1-2 H. L. Clark, 1938
P. inornata 5 4 I A. A. Livingstone, 1933
P. mimica 5 5 1 ” » ”
P. calcar 8 3-4 1 Tasmanian Museum collections
P. gunnii 6 5-6 2 » ” np
P. brevispina .. 6 4-5 2 H. L. Clark, 1938%
* At a size of 10-11 mm. R.
+ H. L. Clark’s deseription mentions the close resemblance of P. regularis to P. nigra but
gives little quantitative data about the latter species.
t See A. M. Clark, 1966, for comments on the specific characters of P. gunnii and P. brevispina,
in Tasmania P. regularis, P. gunnu (Gray), 1840 and P. calcar (Lamarck),
1816 possess gonopores opening abactinally and free swimming larvae; in
P. exigua the gonopore opens actinally and the larvae undergo a shortened
development and in P. vivipara development is internal, there is no gonoduct
and the young emerge in the shear-plane of the interradial are (see Nichols,
1962).
Colour in life may be used to separate the species in the field. P. vivipara
is constantly orange/yellow; in P. exigua the actinal surface is blue-green
and that of P. regularis off-white. P. calcar is noted for its wide range of
colour and pattern and reddish-purple, six armed forms are attributed to
P. gunnii (A. M. Clark 1966).
Acknowledgements
My thanks are due to Miss E. C. Pope and Dr. E. R. Guiler for advice
and criticism. Rererences
BooLooTiaAN, R. A., 1966.—Reproductive Physiology. In ‘Physiology of Echinodermata”,
Boolootian Hid., 598-605. John Wiley, New York.
CuAaRK, A. M., 1966.—Echinodermata. Port Philip Survey, 1957-63. Mem. nat. Mus. Vict.,
27: 320.
CLARK, H. L., 1938.—Echinoderms from Australia. Mem. Mus. comp. Zool. Harvard,
55: 166-169.
, 1946.—The Echinoderm fauna of Australia. Publ. Carneg. Instn. Washington,
U.S8.A., 566: 128-144.
LivinestTong, A. A., 1933—Some genera and species of the Asterinidae. Rec. Aust. Mus.
Sydney, 19: 1-20.
Nicos, D., 1962.—“‘Echinoderms.” 91. Hutchinson University Library. London. 2nd
Impression (revised).
EXPLANATION OF PLATE XXIX
P. vivipara, sp. nov. a. Actinal surface (H372). b. Abactinal surface (H372).
(Photo. C. G. Harrison.) c¢. A specimen with the actinal surface removed. - The sacs
containing the young and developing embryos are indicated. d. Detail of oral and
adambulacral spinulation (H372). e and f. Young emerging through the abactinal
surface of adult (H404).
»
Proc. Linn. Soc. N.S.W., Vol. 98, Part 3 PLATH XXIX
Patiriella vivipara, sp. nov.
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fi
air |<}
Gi iacin echvanou Nona
THE NASAL MITES OF QUEENSLAND BIRDS
(ACARI: DERMANYSSIDAE, EREYNETIDAE, AND
EPIDERMOPTIDAE)
‘Rosert Domrow
Queensland Institute of Medical Research, Brisbane
(Plates xxx-xxx1)
[Read September 25, 1968]
Synopsis
Native and introduced birds of 211 genera and 330 species have been examined
for nasal mites in Queensland. At a generic level, this represents 64% of the Australian,
and 75% of the Queensland fauna. At a specific level, the corresponding figures are
45% and 58%. Of the Queensland fauna, 53% of the genera and 36% of the species
were found to be infested. Three families of mites were discovered in this microhabitat:
Dermanyssidae (eight genera, 99 species), Hreynetidae (four genera, 19 species), and
Epidermoptidae (six genera, six species). Keys are provided to all genera and species
known to occur in Australian birds.
The following 12 dermanyssid species have been described as new: Tinaminyssus
megaloprepiae from Megaloprepia magnifica (Columbidae), TJ. myristicivorae and
T. welchi from Myristicivora spilorrhoa (Columbidae), Ruandanyssus artami from
Artamus cinereus and A. minor (Artamidae), Ptilonyssus neochmiae from Neochmia
phaeton (Ploceidae), P. struthideae from Struthidea cinerea (Corvidae), P. corcoracis
from Corcorax melanorhamphus (Corvidae), P. orthonychus from Orthonyx temminckii
(Timaliidae), P. monarchae from Monarcha trivirgata (Muscicapidae), P. setosae from
Rhipidura setosa and R. rufifrons (Muscicapidae), P. gerygonae from Gerygone
palpebrosa (Sylviidae), and Sternostoma neosittae from Neositta striata (Sittidae).
The following three ereynetid species have been described as new: Boydaia
myzomelae from Myzomela sanguinolenta (Meliphagidae), B. maluri from Malurus
amabilis and M. melanocephalus (Sylviidae), and Ophthalmognathus accipitris from
Accipiter fasciatus (Accipitridae).
Fourteen dermanyssid, eight ereynetid, and one epidermoptid species have been
recorded from Australia for the first time, as have the genera Rhinoecius (Der-
manyssidae), Ophthalmognathus (Hreynetidae), and Mycteroptes (Epidermoptidae).
Forty-nine new host-records have been detailed for 24 dermanyssid species, and
one each for one ereynetid and one epidermoptid species.
Forty generic and 56 specific names (of which 25 and 36, respectively, are new
synonyms) have been discarded as being based on too typological an approach to
taxonomy, without due regard to intraspecific variation and zoogeography.
Five habitats have been sampled more or less intensively, viz. rain-forest, tropical
woodland, tropical tussock grassland, semi-arid savannah, and coastland.
Host-specificity is well marked, each genus or species of mite being restricted
to a particular family, genus, or species of bird. This is true both of cosmopolitan
and Old World groups of birds that occur in Australia (even if represented by only
one species), and of birds such as the honeyeaters (Meliphagidae) that have radiated
in, and are restricted to the Australian zoogeographical region.
The rhinonyssine genera (Dermanyssidae) are considered monophyletic and
separable only on ecological grounds from the ectoparasitic macronyssine species found
on both birds and mammals. In both subfamilies, the protonymph has well developed
chelicerae like those of the adult female, while the deutonymph, with poorly developed
chelicerae like those of the larva, is a non-feeding stage. The genera have developed
along three lines. Tinaminyssus, a simple genus retaining extensive dorsal shields
and a tritosternum in species parasitizing ground-birds, leads, by reductions of the
dorsal armature and the respiratory apparatus, to three genera restricted to birds
of the shore and open water (Larinyssus, Rallinyssus, and Rhinonyssus). All four
possess normal chelicerae with two unreduced digits. In Ruandényssus and Rhinoecius,
the fixed digit is absent, leaving a falcate movable digit of normal proportions. In
Ptilonyssus and Sternostoma, both digits are considerably reduced, with a concomitant
tapering of the distal half of the cheliceral shaft.
PROCEEDINGS OF THE LINNEAN SOcIETY OF NEW SouTH WALES, VoL. 93, Part 3
298 THE NASAL MITES OF QUEENSLAND BIRDS
CONTENTS Page
1. Introduction) 7.) : lke RUIN cases, Gaal my liga Un eaPinint ny MEO eine)
II. Localities and mater Lad tetra iter nero ie an AE)
LM Methods and “materials i. 00 5k ened ee
IV. Systematics os Sein aig nell Crgoh. NG Goel Re eS Oe
V. Host-specificity and pooseetaahs® Vie fo ohl ues. aes |e) toy
Viv Phylogeny of the rhinonyssine genera "2 (02. ee eee
Vil. ‘Host:parasite list “2 (namie eee) ee
I. INTRODUCTION
The first nasal mites of birds were described, sporadically, by European
workers (Giebel, 1871; Berlese and Trouessart, 1889; Tragardh, 1904; Hirst,
19210, b, 1923), and this work was reveiwed and extended by Vitzthum (1935).
More recently, considerable work in Brazil by de Castro (1948), Pereira
and de Castro (1949), and do Amaral (1962 et seq.) has been supplemented
by studies in the U.S.A. by Furman (1957), Strandtmann (1951 et seq.),
Hyland (1961), and Clark (1958 et seq.). Fain’s reviews of the Rhinonyssinae
(1957e), Speleognathinae (1963c), and Turbinoptinae (1960c) are based on
his numerous papers, particularly on the African and European fauna, but
augmented by material from birds of all kinds dying at the Antwerp Zoo.
Zumpt and Till (1955) have also studied the African fauna, Bregetova
(1964 et seq.) has initiated useful studies in the U.S.S.R., and Wilson (1964
et seq.) is working up extensive collections from S.E. Asia and New Guinea.
Lesser papers by several other workers are mentioned whenever necessary
in the text below, and reveiws have been published by Hyland (1963) and
Fain (1965a@). Ticks and chiggers (Acari: Argasidae and Trombiculidae)
have each been once recorded from similar microhabitats in birds (Amerson,
1966; Brennan, 1965), but are not considered below. Another oddity is
the single specimen of Halarachne miroungae Ferris, specific for seals
(Pinnipedia : Phocidae), found in the larynx of a gentoo penguin, Pygoscelis
papua Forster, kept in the same enclosure (Fain and Mortelmans, 1959;
Domrow, 1962).
Ptilonyssus trouessarti Hirst (19210) was the only species known from
Australia when the present study was commenced five years ago. At first,
I published small instalments as material came to hand (Domrow, 1964a
et seq.), but have, for the past three years, determined to assemble material
until the law of diminishing returns had run its course. This last instalment
therefore comprises the briefest possible summary of previous records, as
well as details of all new material collected. Its text was closed on June 30,
1968. An attempt has also been made to list all mite species, which, while
not yet collected, can be expected to occur, together with their hosts, in
Australia. A complete host-parasite list is also given, together with an
indication of the groups of birds that have not yet been examined (Tables
2 and 3), that future collecting, both in Queensland and elsewhere in
Australia, may be directed towards the birds (and habitats) most likely
to prove rewarding. Thus, although no swifts (Apodiformes : Apodidae)
have been examined, this group is known to harbour three rhinonyssines
overseas (Fain, 1957e; Sakakibara, 1967).
All holotypes have been deposited in the Australian National Insect
Collection, C.S.I.R.O. (Canberra), and paratypes, when available, have been
lodged in this Institute, the Prince Leopold Institute of Tropical Medicine
(Antwerp : Dr. A. Fain), and the Bernice P. Bishop Museum (Honolulu :
Dr. R. W. Strandtmann).
ROBERT DOMROW 299
The initials of the collectors, apart from my own, are those of G. J.
Barrow, H. Battam, E. H. Derrick, I. D. Fanning, M. L. Friel (née Emanuel),
B. H. Kay, H. J. Lavery, H. I. McDonald, D. J. Moss, J. Nielsen, H. A.
Standfast, J. S. Welch, and R. H. Whitehead.
II. LocaLitirs AND HABITATS
The localities in Queensland mentioned in Section IV below are indicated
in Fig. la, and have been divided into five major categories, with vegetational
notes after the “Atlas of Australian Resources” (Department of National
Development : Canberra, 1955). Of the New South Wales localities, the
Tweed River flows through an area of subtropical rain-forest, while Urben-
ville is in sclerophyll, and Mt. Keira (near Wollongong) in wet sclerophyll
forest.
(a) Rain-forest—Of the northern localities, the tropical rain-forest
studied at Maalan and Jordan Ck. (both on the escarpment of the Atherton
Tableland near Millaa Millaa) have been discussed more fully by Domrow
(19676). The rain-forest at Ella Bay reaches right to the beach, and is
relatively undisturbed, but that around Innisfail has been largely destroyed
as a result of extensive sugar-cane farming. The Mt. Jukes locality has also
been detailed by Domrow (1967b).
Webb (1959) has pointed out a difference in the rain-forest environment
to the south of Sarina, and the three submontane areas studied in S. E.
Queensland are therefore better classified as subtropical rain-forest. These
are Mt. Glorious, Upper Brookfield, and Wilson’s Peak (Plate xxx, Fig. 1).
(6) Tropical woodland.—Apart from Chillagoe and Charters Towers,
in relatively drier areas of north Queensland, the remaining localities in
this category are in the far south-east of the state (Esk, Cobble Ck., Mt.
Nebo, Samford, Brisbane, Mt. Cotton, Logan Village, Tamborine Village,
Oxenford, and Palen Ck.). These are all in open eucalypt forest, but, in
certain valleys with higher precipitation and denser vegetation, birds typical
of rain-forest, e.g. Macropygia phasianella (Temminck) and Chibia bracteata
(Gould), are commonly seen. The Esk area (Plate xxx, Fig. 2) will be
treated in more detail elsewhere (Domrow, 1969).
(c) Tropical tussock grassland.—This category, mixed with tropical
woodland, is typified by Kowanyama (fomerly the Mitchell River Mission),
and has been treated in detail by Doherty et al. (1963), Standfast (1965),
and Domrow (1967). Viruses isolated from wild birds at this locality have
been detailed by Whitehead et al. (1968). Longreach Lagoon lies between
Gamboola and Highbury cattle stations.
(d) Semi-arid savannah.—Condamine is in an area of tropical woodland
mixed with layered scrub, while Mitchell, Charleville, and Augathella are
in an extensive area of tropical woodland mixed with arid scrub. To the
south, Cunnamulla is surrounded by semi-arid, low-tree savannah, and, further
inland, Winbin Ck. lies in mixed semi-arid shrub savannah and arid. scrub.
Finally, Windorah, in semi-arid tussock grassland, sees the beginning of the
red dunes of the inland desert (Plate xxx1, Fig. 1).
(e) Coastland.—The estuary of Topsy Ck. and Half Tide are vpen ocean
beaches with large areas of sand. Tin Can Bay is a sheltered, very shallow inlet
with a narrow sandy beach above extensive mudflats, the latter being exposed
only at low tide (Plate xxx1, Fig. 2). Cowley Beach and Chelona are examples
_ of mixed coastal woodland, characterized by low forest on impoverished soil,
with extensive areas of mangroves along the water margins.
3500 THE NASAL MITES OF QUEENSLAND BIRDS
Mt Glorious, Cobble Ck
Mt Nebo? *Samford
Upper Brookfield® BRISBANE
Mt Cotton?
Logan Village,
Tamborine Villages .
Oxenford
557
Nem ZA
| ePalen Ck /~
Topsy CkgeKowanyama ie asl ae ered
re 50 miles |
C110
eLongreach_Lagoon
110
Chillagoee
Maalan, | i
Jordan Ck {Cowley Beach
eCharters lowers
%
t Jukes
helona
Half Tide
55
ove %
NOs 2345S
\\
Are
Windorahe
eAugathella
Charleville,
Winbin Cke \ amtchet Tin Can a
Condaminee
eCunnamulla
Wilsons Pk,
150 E “9g
Fig. la. Map of Queensland showing localities mentioned in text (isohyets in
inches/year). -
ROBERT DOMROW 301
IIL. Metruops anp MATERIALS
‘The classical method of collection of intranasal mites is that of Fain
(1957e), and I can do no better than repeat his essentials: “La récolte des
Acariens a été pratiquée généralement peu de temps apres la mort de
VOiseau ... le bee est largement ouvert. Au moyen Vune pure de ciseaua
a mors fins on découpe le palais, le plus pres possible du bec et sur une
grande longueur de fagon @ bien exposer la région qui correspond aux narines.
Les tissus excisés sont ... examinés ultérieurement ... Pour examiner les
narines par Vintérieur il est souvent nécessaire de disséquer les cornets
situés profondément ou les lamelles cornées qui cachent plus ou moins leur
orifice interne.
LAS Epidenmop aides sont généralement cantonnés a Vintérieur méme des
narines et ce n’est qu’en cas de forte infestation qwon peut les voir envahir
la profondeur des fosses nasales. A cause de leur petite taille ils sont souvent
difficiles a découvrir et il est nécessaire Wutiliser un grossissement déja assez
puissant... Les Rhinonyssidés vivent également dans les narines, on peut
cependant les rencontrer aussi dans le mucus qui recouvre les cornets, mais
ils ne s’aventurent jamais trés loin a Vintérieur des fosses nasales. Les
Epidermoptidés comme les Rhinonyssidés sont animés de mouvements lents
et ne se déplacent probablement que trés peu. Il n’en est pas de méme des
Speleognathes qui sont trés rapides et circulent dans toute Vétendue des
fosses nasales, envahissant méme les cellules osseuses”.
IT would only add that I have supplemented this palatal approach with a
dissection inwards and backwards from the nostrils sufficiently deep to expose
the air-sacs of the skull, thus facilitating the collection of speleognathines.
Even this, if neatly performed, does not preclude the preparation of the
skins for museum purposes, and, in any case, the viscera of many of the
specimens were taken for virological and parasitological studies. Both Fain
(1965a) and I (1967a) have commented on the fact that nasal mites are not
found, except for Sternostoma tracheacolum Wawrence and _ occasional
stragglers, in the internal respiratory tract. The work of Maa and Kuo
(1965) would confirm this.
The above method allows the collection of virtually every mite present in
the nasal passages of each bird, and therefore provides a basis for quantitative -
population studies. One such study (Domrow, 1967a) showed no significant
difference in rhinonyssine populations during the wet (March-April) and
dry (October-November) seasons at Kowanyama. However, in the present
study, only one other locality (Esk) was visited regularly, and work here
was directed rather towards a thorough sampling of the bird fauna.
Consequently, once a species was found and confirmed to harbour nasal mites,
it was not collected intensively or ignored at later visits. Further quantitative
data on populations have been supplied by ter Bush (1963) and Amerson
(1967).
The material was fixed in cold 70% ethyl alcohol containing 5%
glycerine, and cleared within a month in warm lactic acid, the latter
procedure causing the legs to extend conveniently (excessive heating causes
cuticular exfoliation, but this is often of little consequence as the setal
bases stand firm). The specimens were then mounted singly (or in pairs if
belonging to a common species) in Hoyer’s formula for gum-chloral medium
(distilled water 50 cc., gum arabic 30 g., chloral hydrate 200 g., glycerine
20 cc., mixed in that sequence at room temperature), and the slides dried
in an oven at 37°C. Hoyer’s, a water-soluble medium, has many advantages
302 THE NASAL MITES OF QUEENSLAND BIRDS
over polyvinyl alcohol, which becomes virtually insoluble and contracts to an
extreme degree with age. I have used Hoyer’s routinely since 1959, and
in the warm-temperate climate of Brisbane, even with its rather humid
summers, no crystallization or blackening has taken place, and ringing
the coverslip with varnish is not necessary. I fancy the presence of a little
lactic acid renders the preparation less prone to dry out at the edge (see
also Womersley, 1943).
IV. SysTeMatics
In this section, the material collected will be dealt with in systematic
sequence, as detailed in Table 1. The following three sections will deal with
host-specificity and zoogeography, the phylogeny of the rhinonyssine genera,
and host-parasite relationships.
I have accepted Evans and Till’s terminology (1965) for the dorsal
shields of the idiosoma, except that I have preferred ‘“mesonotal shieldlets”
to “mesonotal scutellae” (these authors themselves use the term “shield” for
the larger sclerotizations). Also, in the case of the peritrematalia (a term
dating at least from Vitzthum, 1935), I have called the accompanying sclero-
tization, which invariably lies behind the stigma (and peritreme, which is
reduced if present at all), the poststigmatic, and not the peritrematal shield.
TABLE 1
Systematic synopsis of the mites collected
Genera No. of Genera No. of
Family Represented Species Family Represented Species
Dermanyssidae Tinaminyssus 18 Ereynetidae Boydaa 8
(Rhinonyssinae) Larinyssus 2 (Speleognathinae) Neoboydaia 5
Rallinyssus 4 Ophthalmognathus 3
Rhinonyssus 6 Speleognathopsis 3
Ruandanyssus 2 Epidermoptidae Turbinoptes 1
Rhinoecius 2 (Turbinoptinae) Passerrhinoptes 1
Ptilonyssus 53 Oaleya 1
Sternostoma 12 Schoutedenocoptes 1
Rhinoptes 1
Mycteroptes 1
No rigid nomenclature has been adopted for the dorsal setation, as the
frequent reduction renders it difficult always to identify the setae according
to Lindquist and Evans’ system (1965). Nevertheless, it is recognized that
many of the setae present in the Rhinonyssinae have their counterparts in
this system. Thus, on the podonotal shield, my verticals are surely their 41,
and my anterolaterals the intermediate members of their j and 2 series, while
the commonly seen posterodiscal arch of four setae would be their pairs
j5 and 25. The midposterior pair, which, in Ptilonyssus Berlese and Trouessart,
may or may not be left free in the cuticle depending on the degree of trunca-
tion of the shield, is jg. The five pairs of setae between the posterolateral
angles of the shield and the stigmata are r., 75-6, and 85-6 (Ss; being on or off
the shield depending on the degree of its reduction, and absent in some
species of Sternostoma Berlese and Trouessart). The central eight members
of the transverse band of ten setae immediately behind the mesonotal
shieldlets in Ptilonyssus are the pairs J; 2 and Z;». Finally, the pygidial
setae would be J; or Z;. That these setae of the adult are most easily
recognized from Evans and Till’s figure (1965) of the dorsum of the proto-
nymph of Macronyssus flavus (Kolenati), a macronyssine ectoparasite of
bats, is an indication of the reduction common in this endoparasitic group.
ROBERT DOMROW 303
The terminology used for the structures on the venter is also that of
Evans and Till (1965), except that I have preferred the widely used (and
purely Latin) term “adanal” to the hybrid “paranal”’ for the two setae
flanking the anus. The term “cribrum” for the patch of aciculations behind
the postanal seta also dates at least from Vitzthum (1935). I have not
definitely assigned the setae on the ventral cuticle, but, again, the common
occurrence of two setae immediately behind the genital shield, followed by a
transverse row of about six, mirrors more closely the nymphal than the
adult macronyssine pattern.
The legs have been described according to Evans (1963a@), in which the
tarsal setae designated ad, and pd, reduced and often difficult to see in
the rhinonyssines, are equivalent to the two setae noted, but not included
in the tarsal formulae by Domrow (1966)).
Finally, Evans (19638b) and Evans and Till (1965) have been followed
for the terminology of the capitulum, a term they discard in favour of
“onathosoma”, although, for its basal portion, they use “basis capituli’.
I have also preferred to consider the appendage of the palpal tarsus a claw
rather than an apotele (Camin et al., 1958), and have followed Sellnick (1965,
1967) in preferring “epistome” to “tectum”. The palpal setal formulae
commence with the trochanteral setae, and continue, if possible, to the tibiae;
they include the two dorsodistal sensory rods on the tibiae.
Additional morphological studies have been made on the chelicerae,
the genital aperture, the spermatheca and its ducts, and the coxal and
femoral glands (Fain, 1960e, 1963), f, 1966qa).
The foregoing paragraphs refer essentially to the Rhinonyssinae, but
Fain (19626, 19639, h) has also discussed sexual dimorphism, the invaginated
sensory organ of tibia I, and the solenidia of the Speleognathinae.
Family DERMANYSSIDAE
Subfamily RHINONYSSINAE
This is the classification of Evans and Till (1966), the first serious
attempt in two decades to elucidate the suprageneric taxa of the “dermanys-
soid’” and “laelapoid” complexes.
The initial dichotomies of the following key (those based on the nature
of the chelicerae of the adult female, and leading to couplets 3, 6, and 7)
follow the first of Fain’s classifications (1957a@), in which the Rhinonyssidae,
treated as a full family, were divided into three subfamilies : Rhinonyssinae
Trouessart, Rhinoeciinae Fain, and Ptilonyssinae de Castro. The subfamily
Agapornyssinae Gretillat et al. (1959), based on a synonym of Sternostoma
(v. infra), is a synonym of the Ptilonyssinae. Fain was later (1960d) to
erect a fifth subfamily, Larinyssinae, monotypic for Larinyssus Strandtmann,
which, however, I prefer to leave among his rhinonyssine genera.
Further, in more recent work, at least ten new genus-group taxa have
been raised (Brooks and Strandtmann, 1960; Fain, 1964d; Fain and Aitken,
1967; Bregetova, 1965a, b, 1967). Some of these, with composite generic
names, cut across Fain’s lines of division. For example, Rhinosterna Fain
and Sternoecius Fain and Aitken each show characters of Sternostoma
(Ptilonyssinae), and of Rhinonyssus Trouessart (Rhinonyssinae) on the one
hand and of Rhinoecius Cooreman (Rhinoeciinae) on the other. Both are
placed as synonyms of Sternostoma below.
304 THE NASAL MITES OF QUEENSLAND BIRDS
Also, it is clear that other of these taxa are synonyms, even though
certain faunas, e.g. the neotropical and oriental, require further study. For
example, T'rochilonyssus Fain and Aitken shows the process on coxae II and
the elongate sternal and genital shields characteristic of many species of
Ptilonyssus, and, apart from the chelicerae and the armature of the stigmata,
keys directly to that genus in Fain’s system of 1960d. P. maluri Domrow
also shows abnormally long cheliceral digits, and the lack of peritremes
is a criterion, which, in the case of Ptilonyssus and Passeronyssus Fain, is
shown below no longer to hold.
Bregetova (1964) favours a diphyletic system, recognizing an older group
(Rhinonyssidae) and a younger, for which she proposes the new family
Neonyssidae.* In one case, the chelicerae of both nymphal stages are said
to be similar to those of the adult. In the other, there is alleged a “marked
difference between the chelicerae of the feeding protonymph and the non-
feeding deutonymph, the chelicerae of the latter reverting to the form in
the non-feeding larva”. (I use Evans and Till’s words of 1965, but should
note they were phrased for a different context.)
However, this system also cuts across Fain’s, placing Rhinoecius and
Sternostoma among the rhinonyssids, and Ruandanyssus Fain among the
neonyssids, and I believe it to be based on incorrect observation. I have
examined both protonymphs and deutonymphs of Tinaminyssus Strandtmann
and Wharton (many species from pigeons, parrots, kingfishers, and herons),
Larinyssus, Rhinonyssus, Ruandanyssus, and Ptilonyssus, one protonymph
of Rhinoecius, and two deutonymphs of Sternostoma (I have no immature
specimens of Rallinyssus Strandtmann), and, in all cases, they confirm Evans
and Till’s findings, even in such genera as Rhinoecius and Ruandanyssus, in
which the chelicerae lack the fixed digit. Mitchell (1963) has also shown
the phenomenon of alternate feeding stages in his detailed morphological
study of Rhinonyssus rhinolethrum Trouessart, while the pioneer work on
this point was that of Strandtmann (1961).
I therefore consider the Rhinonyssinae a monophyletic group of mac-
ronyssine origin (see Section VI below), and recognize no intermediate
grade between subfamily and genus.
Key to Australian genera of RHINONYSSINAE (females) *
* This and the following keys are all based on Australian material only.
iL, Cheliceral shafts of uniform diameter, the digits occupying more than one-
eighth” of “the total: (length) (ee cee RE a 2
Chelicerae attenuate in distal half, the digits normally occupying. less than
one sixteenth of the total length (but one-eighth in Ptilonyssus maluri
Uf
DONUT OW.) i= 29 Sadi ncsievocas oh agewd Spode date eee onase Bote ocean Oe eee
2.1); Both cheliceral digits) present: ig acgus spencer cps cee iate eae ee ie Cee 3
Mixed 7 chelicéralldisit: absent (yi 2 2. cis. sae eho oe ne erate 6
suC2)ine Stigmata switheiperitremes \4456.,4 20a So. eles eha a Ga ee. Sa eee 4
Stigmata without penitnemesiaaesoeneie ole ee ene Rhinonyssus Trouessart
4 (3). Peritrematalia in normal position above coxae III-IV.
Anus normal | Gad: eset vc eens bee @ Olsen tr ACE ane: Ine 5
Peritrematalia situated near caudal extremity of idiosoma. Some species with
delacatemcincgumianal trill eee re eae ee cee Rallinyssus Strandtmann
5 (4). Podonotal shield always entire. Opisthonotal shield entire, fragmentary, or
ADSEM EM pa, ete ote eee oc OES Tinaminyssus Strandtmann and Wharton
All dorsal shields fragmentary .........0..0../0..: Larinyssus Strandtmann
6 (2). Opisthonotal shield and tritosternum present ............ Ruandanyssus Fain
Opisthonotal shield and tritosternum absent ........ Rhinoecius Cooreman
: * Unfortunately, this author, apparently from a lack of literature, uses some genera
in an outmoded sense. In particular, Neonyssus Hirst has been shown by Fain and
Hyland (1962b) to be a synonym of Ptilonyssus, and her Neonyssidae therefore falls
to de Castro’s Ptilonyssinae (1948).
ROBERT DOMROW 305
7 (1). Stigmata normally with peritremes, but, if not, gnathosoma terminal (except
in P. neochmiae, n. sp.) and anal shield ventral, normally formed, and
beanine spostanalesetay .) oases ceo Ptilonyssus Berlese and Trouessart
Stigmata without peritremes. Gnathosoma withdrawn ventrally between
coxae I at least to level of trochantero-femoral articulation of palpi. Anal
shield terminal, usually regressed, and always lacking postanal seta
Ba of Eid io Buen tence eRe Gk Uae i. ...e.-.....Sternostoma Berlese and Trouessart
Genus TINAMINYsSUS Strandtmann and Wharton
Tinaminyssus Strandtmann and Wharton, 1958, Contr. Inst. Acar. Univ.
Md, 4: 161. Type-species Neonyssus (Ptilonyssoides) trappi Pereira and de
Castro, 1949, Archos Inst. biol., S Paulo, 19: 229; do Amaral, 1967, Ibid.,
34: 187. Mesonyssus Fain, 1960, Revue Zool. Bot. afr., 61: 318, 62: 102.
Type-species Neonyssus treronis Fain, 1956, Ibid., 53: 394. New synonymy.
Mesonyssoides Fain and Nadchatram, 1962, Bull. Annls Soc. r. ent. Belg.,
98: 271. Type-species Mesonyssoides malayi Fain and Nadchatram, 1962,
Loe. cit., 272. New synonymy. Mesonyssoides Strandtmann and Clifford,
1962, J. Parasit., 48: 728. Type-species Mesonyssoides ixoreus Strandtmann
and Clifford, 1962, Loc. cit., 723. New synonymy. Psittanyssus Fain, 1963,
Revue Zool. Bot. afr., 68: 69. Typespecies Psittanyssus baforti Fain, 1963,
Loc. cit., 70. New synonymy. Falconyssus Fain, 1966, Revue Zool. Bot. afr.,
74: 85. Type-species Falconyssus elani Fain, 1966, Loc. cit., 86. New
synonymy.
Both Fain (1963c) and do Amaral (1967) consider the two species
originally placed in Tinaminyssus belong to Mesonyssus, thereby contraven-
ing Art. 23, since the former has priority.
I do not believe the mere absence of the opisthonotal shield in falconyssus
justifies its separation from a genus in which this character is notoriously
variable (see couplets 2 and 3 in the following key, which include the three
species known from Australian psittacids). In any case, this shield is
present in 7. milvi (Fain, 1962c), n. comb., and TJ’. epilews (Wilson, 1964),
n. comb., both of which also have falconiform hosts, and Fain himself has
commented (1968e) : “a notre avis le nombre des écussons dorsaua ne devrait
pas é€tre utilisé commé caractére générique dans la famille Rhinonyssidae”.
Key to females of Australian species of TINAMINYSSUS
iL LrAtOSterniumM spresent i ete. eesti Mette: atceeta Nias cat crete aioe coos uous Sere wishes 2
ATPIOSE RMU Pra SemitweaeAy LS F el NS Bere cael SAR Ia Oe LEVEES. Shem eithe eave ueeiniels 4
2 (1). Opisthonotal shield entire ................ kakatuae (Domrow), n. comb.
Onisthonotaltsmicldt notyemeirey. ks ny care ceesiars & mises aie eeieieusie cucenet: 3
3 (2). Anterior fragment of opisthonotal shield entire, bearing several long, sinuous
setae Postanal seta absent ............ aprosmicti (Domrow), n. comb.
Anterior fragment of opisthonotal shield divided, bearing several short, stiff
setae. Postanal seta present .............. trichoglossi (Domrow), n. comb.
Ae ¢) -Opisthonotalashield: presentiquaticeink « eMlenie scucie crs fie « cieai aie aelece eremicie e.elele =i die 5
Opisthonotaltshileldsralbs enmity avs ets eases: so. ciss Sue ee oa eso sicbe- te lo-cercaele oes ALY
ba¢4) oAdanaltse tae mb elimi a musi yy le tesla oe RA Phil 6
Adanal setae in front of anus or at least level with its centre .............. 7
6 (5). Opisthonotal shield subrectangular, with margins distinct. Femur I (1-4/2-1),
II (1-4/1-1), III (1-3/1-1), IV (1-38/1-0). Genua I-III (1-4/2-1), IV
(1-4/1-0). Capitular setae absent ...... belopolskit (Bregetova), n. comb.
Opisthonotal shield subpentagonal, with eroded margins. Femora 8.7.4.4.
Genua 8.6.6.4. Capitular setae present ........ epileus (Wilson), n. comb.
MED) EEUNOMSCLACT TIME ATCO: WOAS Ay ty sheers coc sci ciercon aaleece tugs uel cove, Sis, a colin Museu Nene) Seu sense saeet 8
Some setae, particularly on coxae, strongly inflated basally ...........:.. 15
SE Cie) aOStAN AIRE SEL ay STE SE Mbt ste eat Stns Meus cybvichauargcteh rei bots fol Slots lame orepcnaycniaes MeetcPee- eek Sikes 9
Postanal seta normaly absent, but minute in specimens of 7. geopeliae (Fain)
RO) COME COMET (GLRIUNEIIN) 66 sco obooebo ba doodeon second eea dodo 11
306 THE NASAL MITES OF QUEENSLAND BIRDS
9 (8). Postanal seta at least as long as adanals ........ melloi (de Castro), n. comb
Postanal seta less than half as long as adanals
10 (9). Adanal setae at least as long as anus. Venter of opisthosoma with about
10 pairs of rather short setae. Basitarsi II-IV 3.3.3 ....................
pretest NSY ahath ta Reale tela eR nigh ate sue OO ra dr Naas 3 ocyphabus (Domrow), n. comb.
Adanal setae barely half as long as anus. Venter of opisthosoma with about
20 pairs of quite long setae. Basitarsi II-lV 4.4.4 hirtus (Wilson), n. comb.
11°°(8))., eAdanaliisetae vat, least. as Jon'sas anus) tec. ade ok ao oe ene eae aft;
'Adanal setae considerably shorter than anus ....columbae (Crossley), n. comb
12 (11). Cribrum absent. Basitarsi II-IV at least 4.4.3. Capitular setae absent
Cribrum present. Basitarsi II-IV 3.3.8. Capitular setae present ............ 14
13 (12). First pair of setae between genital and anal shields minute. All coxae unarmed.
Basitarsi II-I[V 4.4.4. All claws subequal ...... ptilinopi (Wilson), n. comb.
All setae between genital and anal shields subequal. Coxae II-III armed with
erescentic bosses. Basitarsi II-IV 4.4.8. Claws I considerably stronger than
TORS UV 6 Fc hcact SER a AAVA ena OR yn CRASS crn Pees a. EWS Ste ears myristicivorae, D. sp.
14 (12). All setae between genital and anal shields subequal. Coxae without rounded
DOSSC Stee? Ser saicut ts Ste wore gies: S eewenewene or rine ico Re aS EEE ASE aes megaloprepiae, n. sp.
First pair of setae between genital and anal shields considerably weaker
than remainder. Coxae II-IV with distinct, rounded bosses ..............
Ee Ons et tat OEP eG Ore OY Sor eB Dane Goo geopeliae (Fain), n. comb.
15 (7). Adanal setae shorter than anus. Only one seta on coxa I inflated basally.
IBasitarsit VISuV 4 :4e4 ee ae eeeme omens cecum Clie phabus (Domrow), n. comb.
Adanal setae at least as long as anus. Both setae on coxa I inflated basally.
Basitarst, DI-DV 04:43) (3.5. BRAS SSA. | eS ee ee 2
16 (15). Opisthonotal shield rectangular, confined entirely to dorsum. Setae on ventral
surface of opisthosoma simple, including first minute pair. Coxae each
with crescentic boss. All claws subequal. Capitular setae present ........
Re oe, FR Se OO eee ny NEES & Dia SO ore Oe aes macropygiae (Wilson), n. com.b
Opisthonotal shield subcircular, encroaching onto venter. Most setae between
genital and anal shields inflated basally. Coxae II-IV each with stout
spur. Claws I stronger than II-IV. Capitular setae absent welchi, n. sp.
17 (4). Four setae in centre of dorsal surface of opisthosoma much larger than
remainder. SI-II set on sternal shield. Postanal seta present............
St grees Od ahs Fh aes eee re ea ary HE: GE a OS halcyonus (Domrow), n. comb.
All setae on dorsal surface of opisthosoma subequal. Only SI set on sternal
shield. Postanal seta absent ................ daceloae (Domrow), n. comb.
TINAMINYSSUS KAKATUAE (Domrow), n. comb.
Mesonyssoides kakatuae Domrow, 1964, J. ent. Soc. Qd,3: 35. Mesonyssus
kakatuae, Domrow, 1966, Proc. Linn. Soc. N.S.W., 90:191; Wilson, 1966,
Pacif. Insects, 8: 759.
Previous records (both Psittacidae, Psittaciformes).—Red-tailed black
cockatoo, Calyptorhynchus banksii (Latham), Kowanyama. Rose-breasted
cockatoo, Kakatoe roseicapilla (Vieillot), Condamine.
New host record.—Cockatiel, Leptolophus hollandicus (Kerr) (Psitta-
cidae), Condamine, 12.1.1966, R. D., D. J. M., and J. S. W. (6 22,2 64).
TINAMINYSSUS APROSMICTI (Domrow), n. comb.
(Figs 1-6, 8-15)
Mesonyssoides aprosmicti Domrow, 1964, J. ent. Soc. Qd, 3: 26. Mes-
onyssus aprosmicti, Wilson, 1968, J. Parasit., 54: 395. Mesonyssoides
platycerci Domrow, 1964, J. ent. Soc. Qd, 3: 27. New synonymy.
Previous records (both Psittacidae, Psittaciformes).—Red-winged parrot,
Aprosmictus erythropterus (Gmelin), Condamine. Also Kowanyama. Pale-
headed rosella, Platycercus adscitus (Latham), Condamine. Also Esk, Charle-
ville, and Longreach Lagoon.
New host records (all Psittacidae).—King-parrot, A. scapularis (Lichten-
stein), Oxenford, 1.vii.1967, B. H. K. (9 9 2,3 6 46,1 deutonymph). Ring-
ROBERT DOMROW 307
neck parrot, Barnardius barnardi (Vigors and Horsfield), Charleville, 19.1
and 1.ii.1967, R. D. and J. S. W. (14 92,2 ¢ 6); Winbin Creek, 20.1.1966,
R. D., D. J. M., and J. S. W. (1 2). Mulga parrot, Psephotus varius Clark,
Charleville, 23.i.1967, R. D. and J.S.W. (8 22, 2 é 4, 1 protonymph).
Red-backed parrot, P. haematonotus (Gould), Orchard Hills, N.S.W., 22.xii.
1968, D. Himsley (1 92). (This specimen was obtained while I was reading
proof, and is taken into account only here and in Table 4. It is typical of
T. platycerci as originally described.)
The original material of the two taxa here combined was separable
according to the following couplet:
ile Anterior’ fragment of posterior dorsal shield with cluster of 10 enormous
setae. Sternal plate poorly defined. Metasternal setae present. From
an Australian parrot (Aprosmictus Gould) .......... aprosmicti Domrow
Anterior fragment of posterior dorsal shield with cluster of 12 enormous
setae. Sternal plate well defined. Metasternal setae absent. From an
Australian parrot ((LIGtycencus NIZOrs)) aes ee eee platycerci Domrow
The central cluster of enlarged mesonotal setae in 17 female paratypes
from A. erythropterus numbers 9 twice, 10 twelve times, and 11 three times,
while 19 additional females from this host at Kowanyama show 8 once, 9
nine times, and 10 nine times. The cluster is not flanked laterally by an
additional pair of strong setae as described for 7. platycerci.
The central cluster is larger in 9 females from A. scapularis (Fig. 1),
comprising 12 setae three times, 13 twice, 14 twice, and 15 twice (2 of the
cluster of 14 in one specimen are not enlarged). Further, these specimens
do show this central cluster flanked exteriorly by a further pair of enlarged
setae, arranged 1.0 once, 1.1 four times, 1.2 three times, and 0.2 once.
In 43 9 2 and 6 ¢ 6 from P. adscitus, the entire complex of enlarged
mesonotal setae numbers 12 in 37 9 @ and 2 ¢ 6. Five 9 2 and 3 $6
show 11 setae, lacking one of the exteriormost pair or one of the central
cluster. One ¢ and 1 2 show 10 setae, the male lacking two setae from the
central cluster, and the female one of the exteriormost pair and one from
the central cluster. In one female, two setae of the central cluster are
completely fused basally, and set in a single correspondingly larger alveolus.
All specimens from B. barnardi lack the outermost pair of the complex,
the number of setae in the central cluster being 10 five times, 9 ten times
(including both males), and 8 once.
The sternal shield is weakly sclerotized and ill-defined in specimens from
both species of Aprosmictus, but dense and sharply delineated in material
from Platycercus and Barnardius Bonaparte. Metasternal setae are present
in all the specimens from A. erythropterus, although one paratype and four
of the later specimens show a metasternal seta on one side only. The
specimens from A. scapularis (Fig. 2) are uniformly 3.3 (except 3.2 once),
as are those from Platycercus and Barnardius (in both of which latter, SI
is borne on the sternal shield).
The two other species of Tinaminyssus recorded from Australian
psittacids by Domrow (1964a) are TJ. trichoglossi from various lorikeets,
and TJ. kakatuae from crested psittacids (cockatoos and the cockatiel).
Considerable clinal variation is allowed below in the former species, and the
material now discussed from members of a third psittacid group, the parrots,
is similarly considered conspecific.
The material from the mulga parrot (Psephotus Gould) typically shows
only eight enlarged dorsal setae in the mesonotal cluster, and has a sternal
shield intermediate between those of 7. aprosmicti and T. platycerci as
308 THE NASAL MITES OF QUEENSLAND BIRDS
Figs 1-6. Tinaminyssus aprosmicti (Domrow) (2 from Aprosmictus scapularis ).—
1-2, Dorsal and ventral views of idiosoma. 3-4, Dorsal and ventral views of leg IV.
5, Deutosternum and hypostome, with typical setation. 6, Ventral view of capitulum,
with atypical hypostomal setation, i.e. with HI absent (left palp in dorsal view).
(Hach division on the scales equals 100u.) :
Fig. 7. Tinaminyssus trichoglossi (Domrow) (@ from Glossopsitta pusilla) —Dorsal
view of idiosoma.
ROBDRT DOMROW 309
originally described, being tapered anteriorly, leaving SI free in the adjacent
cuticle. Further, there is a concentration of the four midposterior setae
on the podonotal shield not shown in the other material, so best the material
be described, but within my broadened concept of 7. aprosmicti.
Female.—A rounded species when engorged, idiosoma 625y long (calcu-
lated from ruptured specimen illustrated). Podonotal shield (Fig. 14) as
long as wide, 231-264 long, 235-259 wide, well defined, subtriangular, but
with rounded corners and irregular posterior margin; surface closely
ieee vt Ge 13
Figs 8-15. Tinaminyssus aprosmicti (Domrow) (from Psephotus varius) — 8-9, Ventral
and dorsal views of leg III of 9. 10, Chelicera of g. 11, Ventral view of capitulum of 92
(left palp in dorsal view). 12, Sternogenital shield of g. 13-14, Ventral and dorsal
views of idiosoma of 2. 15, Dorsal view of idiosoma of protonymph.
310 - THE NASAL MITES OF QUEENSLAND BIRDS
punctate and marked by muscle insertions. Shield with three setae on each
anterolateral margin; also three pairs of setae in mid-longitudinal line, of
which the central pair is set decidedly further back than in remaining
material. Anterior (mesonotal) fragment of opisthonotal shield not as
strongly delimited, almost semicircular behind rectilinear anterior margin ;
with eight (rarely seven) enlarged setae. Posterior (pygidial) fragment of
opisthonotal shield transversely oval; asetose. Stigmata with much abbrevi-
ated peritremes. Dorsal body cuticle with four shieldlets anterolaterally,
and seven pairs of setae (sixth and seventh pairs enlarged and approximated
as in-original illustration of 7. platycerci).
Sternal shield (Fig. 13) well defined, reticulate, and with some punctae;
tapered distinctly between SI and extending back just behind SII. SIJ-III
free in cuticle. Metasternal setae absent. Genital shield ligulate, irregularly
marked, and bearing two setae; associated pores free in cuticle. Anal shield
rounded anteriorly, more heavily sclerotized laterally; cribrum present. Anus
centrally placed, preceded by adanal setae; postanal seta absent. Ventral
cuticle with ten setae arranged 2.4.4.
Legs with setae short and spinose on anterodorsal aspect, but long and
tapering ventrodistally (Figs 8-9). Coxae 2.2.2.1. Trochanter I (0-1/2-0), II
(2-0/2-0), III-IV (1-0/8-0). Femur I (2-4/2-2), II (2-4/1-1), I1I (1-4/1),
IV (1-3/0-0). Genu I (2-4/2-2), II-III (2-4/0-2), IV (2-4/1-1). Tibiae (2-3/2-1).
Tarsi—.18.18.18 (mv present). Ambulacrum I weaker than IJ-IV.
Basis capituli (Fig. 11) normally with two capitular setae, but one
often absent. Several fine deutosternal denticles present. Hypostome -with
three pairs of setae, HI strongest, HIITI weakest. Palpal tarsus obscured
dorsally by tibia; setal formula 0.2.4.7(6), mostly minute, but one or two
on tibia long and slender. Chelicerae 80-85 long, chelate portion occupying
one-quarter of total length. Tritosternum with hyaline edges on base, and
two distally spiculate laciniae.
Male—lIdiosoma 460y long in relatively unengorged specimen. Dorsum
as in female, but shields slightly smaller; podonotal shield 223x223n.
Sternogenital shield (Fig. 12) irregular in outline, bearing SI, but
SIT-III free in cuticle; genital setae on shield (one missing on one side of
one specimen), with accompanying pores on shield in one specimen, off -in
other. Genital aperture in front of SI. Texture of sternal and genital areas
as in female. Remainder of venter as in female.
Legs as in female, but one seta may be lacking on genua-tibiae Or TV:
one specimen also shows femur IV 4.5.
Capitulum as in female, except for spermatodactyl on chelicera (Fig. 10).
Protonymph.—Dorsum (Fig. 15) as in adult, but shields and mesonotal
setae not so strongly developed.
Sternal area with six setae. Ventral cuticle immediately behind coxae
[V probably with six setae arranged 2.?4. Anal shield flanked posteriorly
by about five setae.
Chaetotaxy of coxae and trochanters as in adult, but trochanter IV not
very clear. Femur J as in adult, IJ probably (1-4/1-1), IIT] (1-3/1-0), IV
as in adult on one side, (1-3/1-0) on other. Genu I (1-4/2-1), I-IV (1-4/0-1).
Tibiae (1-3/2-1). Tarsi—17.17.17 (mv absent). Tarsus I essentially as in
adult. ;
Capitulum essentially as in female.
ROBERT DOMROW 311
TINAMINYSSUS TRICHOGLOSSI (Domrow), n. comb.
(Fig. 7)
Mesonyssoides trichoglossi Domrow, 1964, J. ent. Soc. Qd, 8: 29. Mes
onyssus trichoglossi, Wilson, 1964, Pacif. Insects, 6: 378; 1966, [bid., 8: 766:
1968, J. Parasit., 54: 400. Domrow, 1966, Proc, Linn. Soc. N.S.W., 90: 191.
Mesonyssus neopsittaci: Wilson, 1964, Pacif. Imsects, 6:372; 1966, Jbid.,
8: 759; 1968, J. Parasit., 54: 397. New synonymy. Mesonyssus domicellac
Wilson, 1964, Pacif. Insects, 6:375; 1968, J. Parasit., 54:397. New
synonymy. JMZesonyssus chalcopsittae Wilson, 1968, J. Parasit., 54: 395. New
synonymy.
Previous records (all Psittacidae, Psittaciformes).—Rainbow-lorikeet,
Trichoglossus moluccanus (Gmelin), Tamborine and Logan Villages. Also
Esk and Kowanyama. Scaly-breasted lorikeet, 7’. chlorolepidotus (Kiihl),
Tamborine and Logan Villages. Also Esk. Little lorikeet, Glossopsitta
pusilla (Shaw), Esk.
The above four nominal taxa were based on variation in number and
strength of setae, beginning on the dorsal shields and free segments of the
legs, and extending finally onto the coxae and venter. Moreover, host-
specificity is not absolute (Wilson, 19666), and I now consider this variation
clinal, in the direction: 7. neopsittaci—unnamed form from Glossopsitta
(Fig. 7)—typical form of 7. trichoglossi—unnamed form from T'richoglossus
recognized by Domrow (1964) and figured by Wilson (1964)—T. domicellae—
T. chalcopsittae.
The following specimens housed in the Australian Museum, Sydney, and
examined through the courtesy of Mr. C. N. Smithers, are listed for historical
reasons, being among the first nasal mites of birds ever collected. Prior
to 1892, all known species were European, one having been described by
Giebel (1871) and three by Berlese and Trouessart (1889). The first Aus-
tralian rhinonyssine was described by Hirst (19216), and work in this region
was resumed by Domrow (1964a et seq.).
The series comprises 32 2 (in fair condition) from T. moluccanus
(listed on the label under its synonym 7’. multicolor (Gmelin) ), Tweed River,
N.S.W., 11.1892, T. Steel (Carter, 1926, Proc. Linn. Soc. N.S.W., 51: vii.
gives a brief biography). All the specimens agree with the variety noted
in the original description.
TINAMINYSSUS BELOPOLSKII (Bregetova), n. comb.
Neonyssus belopolskti Bregetova, 1950, Dokl. Akad. Nauk SSSR, 71:
1005; Strandtmann, 1956, J. Kans. ent. Soc., 29: 137; Mitchell, 1961, SWest.
Nat., 6: 105. Mesonyssus belopolskii, Domrow, 1965, Acarologia, 7: 433:
1966, Proc. Linn. Soc. N.S.W., 90: 191; Wilson, 1968, J. med. Ent., 5: 213.
Neonyssus ardeae Zumpt and Till, 1955, J. ent. Soc. sth. Afr., 18: 63; Fain,
1957, Annls Mus. r. Congo belge Sér. 8vo, 60:53, 187. Neonyssus bubulci
Zumpt and Till, 1955, J. ent. Soc. sth. Afr., 18: 66; Fain, 1957, Annis Mus.
r. Congo belge Sér. 8vo, 60: 54. New synonymy. Neonyssus ixobrychi Fain,
1956, Revue Zool. Bot. afr., 53:134; 1957, Annls Mus. r. Congo belge Sér.
8vo, 60: 54. New synonymy. Neonyssus marcandrei Gretillat, Capron, and
Brygoo, 1959, Acarologia, 1: 379. Mesonyssus belopolskti nycticoracis Fain,
1961, Acarologia, 3: 516.
Previous records (all. Ardeidae, Ciconiiformes).—Plumed egret, Egretta
intermedia (Wagler). Little egret, EH. garzetta (Linnaeus). Cattle egret
(introduced), Bubuleus ibis (Linneaus). White-faced heron, Notophoys
novaehollandiae (Latham), Esk. Also Charleville, Tin Can Bay, Mt. Jukes,
Slee THE NASAL MITES OF QUEENSLAND BIRDS
and Kowanyama. Pied heron, Notophoysx picata (Gould), Kowanyama. Little
bittern, Ixobrychus minutus (Linnaeus).
I have been able to examine the following material of N. bubulci from
the type-host, B. ibis: 1 @ and 1 ¢ paratype (Transvaal), 1 2? and 1 ¢
(Bechuanaland), and 2 2 @ (Rwanda), the last four specimens bearing
Fain’s identifications. As described, this taxon shows the opisthonotal shield
broader than is usual in the other five taxa listed above, but one of the
two specimens from Rwanda is intermediate in this respect, and I therefore
accept the synonymy. The synonymy of NV. ivobrychi, reasonably clear from
the published data, is confirmed by a study of the holotype @ from Rwanda.
All six taxa are restricted to ardeid hosts.
TINAMINYSSUS EPILEUS (Wilson), n, comb.
Mesonyssus epileus Wilson, 1964, Pacif. Insects, 6: 358; 1965, Ibid.,
7: 638.
Previous record.—Crested hawk, Aviceda subcristata (Gould) (Accipi-
tridae, Falconiformes).
Three paratype @ 2 from the above (type) host all show trochanter
II 5, rather than 4.
TINAMINYSSUS MELLOI (de Castro), n. comb.
Neonyssus (Neonyssus) melloi de Castro, 1948, Archos Inst. biol., S
Paulo, 18: 270. Mesonyssus melloi, Domrow, 1965, Acarologia, 7: 484; 1966,
Proc. Linn. Soc. N.S.W., 90: 192; Wilson, 1966, J. Parasit., 52: 1211.
Mesonyssus melloi streptopeliae Fain, 1962, Revue Zool. Bot. afr., 65: 310.
Mesonyssus streptopeliae, Fain, 1965, Revue Zool. Bot. afr., 72: 159.
Neonyssus hirsutus Feider, 1962, Studii Cerc. stiint. Idsi, 13: 58. New
synonymy.
Previous records (all introduced Columbidae, Columbiformes) .—
Domestic pigeon, Columba livia Gmelin, Brisbane. Indian turtle-dove, Strep-
topelia chinensis (Scopoli), Brisbane. Senegal dove, S. senegalensis
(Linnaeus).
New host records—White-headed pigeon, C. norfolciensis WLatham,
Wilson’s Peak, 15.v.1967, R. D. and B. H. K. (16 @¢@, 11 3 ¢). Wonga
pigeon, Leucosarcia melanoleuca (Ieee) (Columbidae), Mt. Keira, 19.xi.
UWDEG, lel 1B, (bee )).
This latter host was banded by Mr. H. Battam, Cronulla, who noted
several of these mites in and around the right nostril. When they were
disturbed, two or three crawled into the open beak, others withdrew into
the nostril, and only one was captured.
TINAMINYSSUS OCYPHABUS (Domrow), n. comb.
Mesonyssus ocyphabus Domrow, 1965, Acarologia, 7: 487.
Previous record.—Crested pigeon, Ocyphaps lophotes (Temminck)
(Columbidae, Columbiformes), Samford and Condamine. Also Esk, Charle-
ville, Augathella, Longreach Lagoon, and Kowanyama.
New host record.—Squatter-pigeon, Geophaps scripta (Temminck)
(Columbidae), Kowanyama, 1.iv.1965, R. D. (2 2 9).
My original comparison (1965c) of the legs of this species with those
of T. geopeliae Fain (1964c) would imply that ambulacrum I is modified.
In fact, it does not differ appreciably from TI-IV.
ROBDRT DOMROW 313
Both specimens from Geophaps Gray are typical of the original
description, except that the posterior seta on coxae I-III is relatively weaker,
only 6-7 pairs of setae are present on the ventral body cuticle, and the
central two pairs of setae on the opisthonotal shield are obsolescent or
absent. Leg chaetotaxy as in specimens from the type host, 0. lophotes.
Coxae 2.2.2.1. Trochanters I-III (1-0/2-1), IV (1-0/3-1). Femora I-II
(1-4/3-1), III-IV (1-8/1-0): Genu I (1-4/2-1), II-III (1-4/0-1), IV
(1-3/0-0). Tibiae (1-8/2-1). Tarsi—17.17.17 (mw present; basitarsi
trisetose).
TINAMINYSSUS HirRTUS (Wilson), n. comb.
Mesonyssus hirtus Wilson, 1966, J. Parasit., 52: 1210; 1968, J. med. Ent.,
5: 215.
This species, originally described from a member of the genus Chalcophaps
Gould in the Philippines and New Guinea, may now be recorded from
Australia: 4 9 @ from a green-winged pigeon, C. chrysochlora (Wagler)
(Columbidae, Columbiformes), Wilson’s Peak, 15.v.1967, R. D. and B. H. K.
TINAMINYSSUS COLUMBAE (Crossley), n. comb.
Neonyssus (Neonyssus) columbae Crossley, 1950, Proc. ent. Soc. Wash.,
52: 309. Mesonyssus columbae, Domrow, 1965, Acarologia, 7: 440; Wilson,
1966, J. Parasit., 52: 1211.
Previous records (both Columbidae, Columbiformes).—Domestic pigeon
(introduced), Columba livia Gmelin. Crested pigeon, Ocyphaps lophotes
(Temminck), Esk.
TINAMINYSSUS MEGALOPREPIAE, N. Sp.
(Figs 16-24)
Diagnosis.—In Wilson’s key (1964) to the species of Tinaminyssus
from columbiforms, 7. megaloprepiae comes nearest to T. ptilinopi (Wilson).
However, the new species differs in showing (i) the opisthosomal setae few
and uniform ventrally, i.e. midanterior pair not obsolescent; (ii) a cribrum;
and (ii) distinct capitular setae and deutosternal denticles. The setal
formulae for the legs given below also differ from those provided by Wilson,
although I should note that, in a paratype from Ptilinopus perlatus zonurus
Salvadori, trochanter I is (1-0/2-1) and the tibiae (1-3/2-1).
Specimens of 7. phassae (Wilson, 1966a), n. comb., lacking the postanal
seta would also key to this couplet, but 7. megaloprepiae is distinct in the
same three items cited above. There are also differences in the setal formulae
of the leg segments, although I should note that a paratype of T. phassae
shows trochanter I (1-0/2-0), genu I (1-4/2-1), and tarsi—.18.18.17 (mv
present, basitarsi II-III with four, but IV with three setae).
Types.—Holotype female, allotype male, two paratype males, and one
protonymph from a wompoo pigeon, Megaloprepia magnifica (Temminck)
(Columbidae, Columbiformes), Maalan, 17.viii.1965, R. D. and J. S. W. Holo-
type and allotype N. I. C.; paratypes R. D.
Female.—Idiosoma 737 long in slightly distorted specimen figured.
Podonotal shield (Fig. 16) 281x304y, well defined, but with irregular margins;
surface reticulate, closely punctate, and marked by muscle insertions; with
13 setae and two pores. Opisthonotal shield 322x317», of similar texture,
narrowed posteriorly, and not encroaching onto venter; with six setae (one
c
314° THE NASAL MITES OF QUEENSLAND BIRDS
of which is double) and 183 pores arranged as figured. Marginal cuticle
with about four pairs of setae. All dorsal setae minute, slightly tapered
rods. Peritremes as long as diameter of stigmata; poststigmatic shields
absent. R
Sternal shield irregular (Fig. 17), delimited only by cessation of
cuticular striae, textures except for few punctae, without pores. SI-III
increasing in length in that order, slightly inflated basally; SI-II sub-
marginal, III free in cuticle. Metasternal setae absent. Genital shield drop-
‘Figs 16-24. Tinaminyssus megaloprepiae, n. sp. (from Megaloprepia magnifica) — 16-17,
Dorsal and ventral views of idiosoma of 9. 18, Ventral shields of ¢. 19, Ventral view
of capitulum of 9, with deutosternum omitted (right palp in dorsal view). 20, Ventral
view of basis capituli of g. 21, Chelicera of ¢.. 22, Dorsal view of tarsus I of 9. 23-24,
Dorsal and ventral views of leg IV of 9.
ROBDRT DOMROW 315
shaped, shagreened, with muscle insertions and rayed operculum. Genital
setae absent, but attendant pores present in adjacent cuticle. Anal shield
almost twice as long as wide (1384x76,), rounded anteriorly, and with sub-
parallel, sclerotized lateral margins. Definite cribrum present. Anus centrally
placed, preceded by elongate adanal setae; postanal seta absent. Ventral
cuticle with several pores and 12 basally expanded setae, of which mid-
anterior two are not at all reduced, although posterior five are.
Legs with setae on coxae and trochanters similar to those on venter,
but remainder generally much shorter and more spinose, particularly on
dorsal aspect (Figs 23-24). Coxae without ventral excrescences. Coxae
2.2.2.1. Trochanter I (1-0/2-1), II (2-0/2-1), ITI-IV (2-0/2-0). Femur I
(1-4/2-1), II (1-4/1-1) on one side and (1-4/2-1) on other, rae (1- ch —0),
IV (1-3/1-0). Genu I (1-4/2-1), II-IV (1-4/0-1). Tibiae (1-8/2-1). Tarsi -.
17.17.17 (mv present, but basitarsi trisetose). Ambulacrum I with pretarsus
much shorter, claws slightly stronger (and with correspondingly larger
articulatory sclerites), and pulvillus less extensive than II-IV (Fig. 22).
Basis capituli with two capitular setae (Fig. 19); deutosternal denticles
as in male (Fig. 20). Hypostome with three pairs of setae. Palpal tarsus
not completely obscured dorsally by tibia; setal formula 1.3.3/2.8 (vague,
apparently internal structure figured between two ventrodistal setae of
tibia may be remnant of claw). All setae on capitulum in form of stout
rods, except those on more distal segments of palpi (especially one on tarsus).
Chelicerae not in full lateral view, but 116» long, with chelate portion
occupying one-quarter of total length. Tritosternum absent.
Male.—Idiosoma 550-605, long in slightly distorted specimens. Dorsum
as in female, but with some tendency for opisthonotal shield to encroach
onto sides of body. Podonotal shield 250-264, long, 273-286» wide.
Venter as in female, except for genital aperture in front of SI (Fig. 18).
Legs as in female, but femur I (1-4/1-1) once, II (1-4/2-1) twice, IIT
(1-4/0- 0) twice and (1-3/0-0 ) twice ; oon IV (1-38/0-1) once. One specimen
is teratological, showing femur tibia I (1-0/2-1) because all 4.4.3 setae on
dorsal aspect are lacking.
Capitulum as in female, with distinct deutosternal denticles (Fig. 20).
Chelicerae. with spermatodactyl (Fig. 21).
TINAMINYSSUS GEOPELIAE (Fain), n. comb.
Mesonyssus geopeliae Fain, 1964, Revue Zool. Bot. afr., 70: 33; Domrow,
1965, Acarologia, 7: 440; 1966, Proc. Linn. Soc. N.S.W., 90: 192; Wilson,
1966, J. Parasit., 52: 1211.
Previous records (both Columbidae, Columbiformes).—Peaceful dove,
Geopelia placida Gould, Kowanyama. Also Charleville. Bar-shouldered dove,
G. humeralis (Temminck), Samford. Also Esk and Kowanyama.
New host record.—Diamond-dove, G. cuneata (Latham), Charleville, 1.ii-
1967, R. D. and J. 8S. W. (6 2 9, Aen)
The specimens from G. cuneata closely resemble the original descriptions,
except that a minute postanal seta is present, and the seta in the posterior
angles of coxae J-IIT, and the bosses on coxae II-IV are weaker.
TINAMINYSSUS PTILINOPI (Wilson), n. comb.
Mesonyssus ptilinopi Wilson, 1964, Pacif. Insects, 6: 366; 1965, Ibid.,
7: 638; 1966, J. Parasit., 52: 1212; 1968, J. med. Ent., 5: 221.
Previous record. —Purple- crowned pigeon, Ptilinopus superbus (Tem-
minck) (Columbidae, Columbiformes).
316° THE NASAL MITES OF QUEENSLAND BIRDS
TINAMINYSSUS MYRISTICIVORAR, 0. Sp.
(Figs 25-33)
Mesonyssus sp. “A” Domrow, 1967, Proc. Linn. Soc. N.S.W., 91: 215.
Diagnosis—In Wilson’s key (1964) to the species of Tinaminyssus
from columbiform birds, 7. myristicivorae comes nearest T. treronis (Fain),
n. comb. However, the new species differs in showing (7) considerably shorter
setae ventrally on the legs and opisthosoma, on which latter the midanterior
Figs 25-33. Tinaminyssus myristicivorae, n. sp. (2 from Myristicivora spilorrhoa) —
25-26, Dorsal and ventral views of idiosoma (opisthonotal shield incomplete). 27,
Opisthonotal shield (much flattened). 28, Anal shield. 29-30, Dorsal and ventral views
of tarsus I. 31-32, Dorsal and ventral views of leg IV. 33, Ventral view of capitulum
(right palp in dorsal view).
ROBPRT DOMROW S17
pair is not obsolescent; (7) claw I considerably stronger than TI-IV; and
(ui) no elongate seta on the palpal tibiotarsus.
There are also differences in leg chaetotaxy, that of the holotype of
T,. treronis being coxae 2.2.2.1; trochanter I (0-1/2-1), IIT (0-0/3-1), III
(1-0/3-0), IV (1-0/4-0) ; femur I (0-4/3-1), IT (0-4/2-1), III (0-4/2-0),
IV (0-3/2-0); genu I (1-4/2-1), II-III (0-4/2-1), IV (0-4/2-0); tibiae
(1-8/2-1) ; tarsi—.17.17.17 (mv present, but basitarsi trisetose).
T. myristicivorae is separable from the more recently described 7’. phassae
(Wilson, 1966a) in showing considerably enlarged claws on tarsi I and
uniform ventral opisthosomal setae. See also above diagnosis of YT.
megaloprepiae, n. sp., for comments on the leg setation of 7’. phassae, which
also differs in detail from that of 7. myristicworae.
Types—Holotype female and two paratype females from a Torres Strait
pigeon, Myristicivora spilorrhoa (Gray) (Columbidae, Columbiformes),
Kowanyama, 22.x.1966, R. D. Holotype N. I. C.; paratypes R. D.
Female.—Idiosoma 605-638. long in ruptured specimens. Podonotal
shield (Fig. 25) 273-276 long and 366-370” wide; well defined anteriorly
and posteriorly, but with eroded margins and posterolateral excavations at
level of peritremes. Surface reticulate, closely punctate, and marked by
muscle insertions; with eight pairs of minute setae and three pairs of pores.
Opisthonotal shield (Fig. 27) of similar texture, expanded posterolaterally
to encroach onto venter; with four or five pairs of minute setae and about
eight pairs of pores arranged as figured. Marginal cuticle with few pores
near peritremes. Poststigmatic shields absent.
Sternal shield (Fig. 26) indicated only by cessation of cuticular striae;
without pores, but with few scale-like markings. SI submarginal, but II-III
free in adjacent cuticle. Metasternal setae absent. Genital complex as in
T. megaloprepiae, n. sp. Anal shield (Fig. 28) slightly longer than wide,
102x94, more heavily sclerotized marginally; without cribrum. Anus
centrally placed, with adanal setae near its anterior margin; postanal seta
absent. Ventral cuticle with about five pairs of short setae, which are only
slightly expanded basally, and of which midanterior pair is not obsolescent.
Leg setae also short, particularly on dorsal surface (Figs 31-32). Coxae
II-III with sclerotized crescent posteroventrally. Coxae 2.2.2.1. Trochanter
I (1-0/2-1), IT (2-0/2-1), ITI-IV (2-0/2-0). Femur JI (1-4/3-1) but
(1-4/2-1) twice, II (1-4/2-1), TIT (1-4/1-1) but (1-4/2-1) once, IV
(1-4/1-0). Genu TI (1-4/2-1), II-III (1-4/0-1), IV (1-4/1-1). Tibiae
(1-3/2-1). Tarsi—.18.18.17 (mv present; basitarsi II-III with four, but IV
with three setae). Ambulacrum I (Figs 29-30) with very short pretarsus
and internally sclerotized, truncate pulvillus. Claw I very considerably
stronger than IT-IV.
Basis capituli (Fig. 33) without capitular setae and deutosternal
denticles. Hypostome with three pairs of subequal, spinose setae. Palpal
tarsus completely obscured dorsally by tibia; setal formula 0.1.3.6. Chelicerae
116 long, with chelate portion occupying one-quarter of total length.
Tritosternum absent.
TINAMINYSSUS PHABUS (Domrow), n. comb.
Mesonyssus phabus Domrow, 1965, Acarologia, 7: 435.
Previous record.—Common bronzewing, Phaps chalcoptera (Latham)
(Columbidae, Columbiformes), Esk. Also Charleville.
318 THE NASAL MITES OF QUEENSLAND BIRDS
' Figs 34-38. Tinaminyssus macropygiae (Wilson) (from Macropygia phasianella) —34—85,
Dorsal and ventral views of idiosoma of 9. 36, Ventral shields of g. 37, Ventral view
of capitulum of 9, with dorsal setation of right palp not shown. 38, Ventral view of
leg I of 9.
ROBDRT DOMROW 319
TINAMINYSSUS MACROPYGIAH (Wilson), n. comb.
(igs 34-38)
' Mesonyssus macropygiae Wilson, 1966, Pacif. Insects, 8: 607; 1968, J.
med. Hnt., 5: 215.
This species may now be recorded from Australia, the original series
being from New Guinea and the Philippines: 19 9 9 and 4 ¢ 4 from brown
pigeons, Macropygia phasianella (Temminck) (Columbidae, Columbiformes),
Ella Bay and Jordan Creek, 4 and 18.viii.1965, respectively, R. D. and J. 8. W.;
also 10 2 2 and 2 ¢ 6, Wilson’s Peak, 15.v.1967, R. D. and B. H. K.
Dr. Wilson has kindly confirmed the identity of my material.
TINAMINYSSUS WELCHI, 0. Sp.
(Figs 39-47)
Mesonyssus sp. “B” Domrow, 1967, Proc. Linn. Soc. N.S.W., 91: 215.
Diagnosis—In Wilson’s key (1964), 7. welch falls near TJ. gourae
(Wilson), n. comb. The two species also show ambulacrum I modified and
(as far as can be judged from Wilson’s heavily stippled figure) podonotal
and opisthonotal shields similar in both their shape and their setation.
However, the basally expanded setae on all coxae and the venter of 7’. welch
immediately separate the two species, and further minor differences are to
be seen in. the armature of coxa IV and the width of the genital shield.
Of the more recently described species from the Australian region,
T. phabus (Domrow, 1965c) and T. macropygiae (Wilson, 1966a@) also show
coxal processes and basally expanded setae on the legs, but differ from
T. welchi in the details of this armature as well as in the shape and setation
of the opisthonotal shield and the setation of the venter.
Of the remaining two parasites of columbids briefly diagnosed by Wilson
(1966a, 19686), 7. angustus (Wilson), n. comb., is said to have a rectangular
opisthonotal shield, and 7’. peleiae (Wilson), n. comb., distinct poststigmatic
shields.
Finally, 7. welchi recalls the neotropical 7. castroae (do Amaral, 1963q@),
n. comb., which is, however, unique in showing basally expanded genital and
sternal setae as well.
Types.—Holotype female, allotype male, five paratype females, and one
deutonymph from a Torres Strait pigeon, Myristicivora spilorrhoa (Gray)
(Columbidae, Columbiformes), Kowanyama, 22.x.1966, R. D. Holotype and
allotype N. I. C.; paratypes R. D. and A. F.
Female.—Idiosoma (moderately engorged) 627-660» long and 385-396
wide in four somewhat compressed specimens; 627 x 374y in one unflattened
specimen. Podonotal shield (Fig. 39) irregularly arched anteriorly, excavated
posterolaterally around stigmata, and rectilinear posteriorly ; with eight pairs
of minute setae in addition to paired pores and muscle insertions. Opis-
thonotal shield concave anteriorly and expanded posterolaterally to encroach
broadly (Fig. 41) onto venter in unfed, and slightly so in fed specimens;
with four pairs of setae (including pygidial pair) in addition to paired
pores and muscle insertions. Both shields distinctly reticulate and heavily
stippled in entirety. Stigmata with short peritremes; poststigmatic shields
absent; adjacent cuticle with four pairs of minute setae.
Sternal shield (Fig. 40) subpentagonal, delineated only by cessation of
cuticular striae, very weakly sclerotized, and textureless except for weak
scalation; SI and II on shield, III free in cuticle (only one SI in one speci-
men). Metasternal setae absent. Genital shield narrow, shagreened, and with
320 THE NASAL MITES OF QUEENSLAND BIRDS
‘Figs 39-47. Tinaminyssus welchi, n. sp. (from Myristicivora spilorrhoa) —39—40, Dorsal
and ventral views of idiosoma of 9. 41, Opisthonotal shield of unengorged 9. 42,
Sternogenital shield of g. 43-44, Dorsal and ventral views of leg IV of 9. 45-46, Dorsal
and ventral views of tarsus I of 9. 47, Ventral view of capitulum of 9, with inset of
chelicera (left palp in dorsal view).
ROBERT DOMROW 321
weak muscle insertions; operculum rayed; genital setae as in 7’. megalo-
prepiae, n. sp. Anal shield well sclerotized marginally, but transparent
centrally; adanal setae level with front of anus, subequal to postanal seta;
cribrum present. Ventral cuticle with two to four pores and 34-42 setae,
of which 14-16 are basally expanded, with short terminal filament, while
remainder are normal, 7%. e. elongate.
Coxae IJ-IV armed posteriorly—this armature is not simply the crescentic
thickening found in 7’. ocyphabus (Domrow, 1965c), or the rosette of 7.
geopeliae (Fain, 1964c), but a strong, distinctly protruding spur analogous
to those present in many species of Hchinonyssus Hirst and T'richosurolaclaps
Womersley (see Domrow, 1966a). Coxae 2.2.2.1. Trochanter I (1-0/2-1),
II (1-0/3-1) but (1-0/3-0) once, III-IV (1-0/3-0). Femur I (1-4/3-I), I
(1-4/2-1) but (1-4/8-1) twice and (1-4/2-0) once, III (1-5/1-0) but (0-5/
1-0) once, IV (1-4/0-0) but (0-4/0-0) once. Genu I (1-4/2-1), IJ-IV
(1-4/0-1). Tibiae (1-8/2-1). Tarsi—.18.18.17 (mv present; basitarsi II-III
with four, but IV with three setae). Only the above six variations were noted
in the 270 segments checked. Both setae on coxa J, anterior seta on coxae II
and III, and seta on coxa IV modified; posterior seta on coxa II and III
normal. Modified setae present on venter of remaining segments as follows: one
to three on dise of trochanters, and one (often weak) on femur-tibia I and
femora and tibiae II-IV. Some tarsal setae slightly expanded basally.
Tarsus I incrassate (igs 45-46), strongly sclerotized, and with dorsodistal
sensory islet; pretarsus shorter, claws stronger, and pulvillus smaller (yet
less diaphanous) than II-IV.
Basis capituli (Fig. 47) well sclerotized; with about six rows of
deutosternal denticles, but without capitular setae. Hypostome with three
pairs of subequal setae. Palpi with five free segments, but tarsus obscured
dorsally by tibia; setal formula 1.1.4.6. Tarsus with about seven minute
setae. All setae on capitulum spine-like. Chelicerae with digits occupying
one-quarter of total length (Fig. 47). Tritosternum absent.
Male.—Similar to female, but smaller, idiosoma 495, long.
Genital area of sternogenital shield hyaline marginally, with very
irregular, but strongly sclerotized patch discally (Fig. 42). Venter of
opisthosoma with four expanded setae on one side and seven on other;
eleven normal setae on each side.
Leg chaetotaxy as in female, except coxa II lacking pv, femur III
(0-5/1-0), and IV (1-4/1-0) (all on one side only).
Spermatodactyl as long as cheliceral digits.
TINAMINYSSUS HALCYONUS (Domrow), n. comb.
Mesonyssus halcyonus Domrow, 1965, Acarologia, 7: 448; Wilson, 1966,
Pacif. Insects, 8: 609; 1968, J. med. Ent., 5: 213. Falconyssus halcyonus,
Fain, 1966, Revue Zool. Bot. afr., 74: 86.
Previous records (both Alcedinidae, Coraciiformes).— Sacred kingfisher,
Halcyon sanctus Vigors and Horsfield, Logan Village and Brisbane. Also
Ksk, Charleville, Winbin Ck., Half Tide, Innisfail, and Kowanyama. Mangrove-
kingfisher, H. chloris (Boddaert).
New host records——Red-backed kingfisher, H. pyrrhopygius Gould,
Kowanyama, 27 and 30.iii, and 17.iv.1965, R. D. and J. S. W. (112 2,24 4,
4 nymphs). Forest kingfisher, H. macleayii Jardine and Selby, Innisfail,
3.vili.1965, R. D. and J. S. W. (222 2, 6¢ ¢, 3 nymphs); Kowanyama,
2931, ( and 14 iv.1965, and 12111-1967, R. D., B. H. K., H. A. 8S. and J.S. W.
(Seo a)
322° THE NASAL MITES OF QUEENSLAND BIRDS
All specimens are similar dorsally. However, those from H. pyrrhopygius
and H. macleayti show pointed, not spatulate tarsal claws, while the setae
on the ventral cuticle are only slightly inflated basaily in the former series,
and barely at all in the latter. ‘
TINAMINYSSUS DACELOAE (Domrow), n. comb.
Mesonyssus daceloae Domrow, 1965, Acarologia, 7: 445. Falconyssus
daceloae, Fain, 1966, Revue Zool. Bot. afr., 74: 87.
Previous records (both Alcedinidae, Coraciiformes).—Laughing kooka-
burra, Dacelo gigas (Boddaert), Samford and Esk. Also Condamine and
Mt. Jukes. Blue-winged kookaburra, D. leachit Vigors and Horsfield, Chelona.
Also Kowanyama.
I regret having formed this specific name in the first, rather than the
third declension—Dacelo Leach is an anagram of Alcedo Linnaeus, and is
the basis of the subfamily Daceloninae (see Thomson, 1964). Art. 32, however,
forbids any change (Stoll et al., 1964).
Genus Larinyssus Strandtmann
Larinyssus Strandtmann, 1948, J. Parasit., 34:507. Type-species
Larinyssus orbicularis Strandtmann, 1948, Loc. cit., 507.
Key to females of Australian species of LARINYSSUS
iL, Ventral surface of opisthosoma with about ten pairs of setae. Capitular
setae present. Two pairs of hypostomal setae present ........ benoiti Fain
Ventral surface of opisthosoma with about three pairs of setae. Capitular
setae absent. Three pairs of hypostomal setae present ................
TT cache 1c oats REE Pee Se eat EES, kre ehawe orbicularis Strandtmann
LARINYSSUS BENOITI Fain
Larinyssus benoiti Fain, 1961, Revue Zool. Bot. afr., 68: 128; 1964,
Annls Mus. r. Afr. cent. Sér. 8vo0, 182: 134; Domrow, 1966, Proc. Linn. Soc.
N.S.W., 90: 196. |
Previous records (both Glareolidae, Charadriiformes).— Australian
pratincole, Stiltia isabella (Vieillot), Kowanyama. Oriental pratincole,
Glareola pratincola (Linnaeus).
Fain’s single female (19600) was originally misidentified as the follow-
ing species (personal communication).
LARINYSSUS ORBICULARIS Strandtmann
Larinyssus orbicularis Strandtmann, 1948, J. Parasit., 34: 507; Zumpt
and Patterson, 1951, J. ent. Soc. sth. Afr., 14:78; Bregetova, 1951, Parazit.
Sb., 138: 117; Fain, 1956, Revue Zool. Bot. afr., 538: 157. Larinyssus petiti
Gretillat, 1961, Vie Milieu, 12: 155. New synonymy.
Previous records (all Laridae, Charadriiformes).—White-winged black
tern, Chlidonias leucoptera (Temminck). Gull-billed tern, Gelochelidon
nilotica (Gmelin). Common tern, Sterna hirundo Linnaeus. Sooty tern,
Sterna fuscata Linneaus. This record is based on host-relationship (Amerson,
1967), and I have been unable to obtain specimens to confirm it. Southern
black-backed gull (vagrant), Larus dominicanus Lichtenstein.
This widespread parasite of gulls and terns may now be recorded from
Australia: 16 2 2, 2 ¢ 3, and 1 protonymph from a crested tern, Sterna bergit
Lichtenstein (Laridae), estuary of Topsy Creek, 29.x.1966, R. D. and H. A. S.
_ Repeated attempts to borrow material of Gretillat’s species have been
In vain, but the host data and some of the original figures make me certain
<s
te
w
ROBERT DOMROW
the synonymy is correct. The alleged single dorsal shield, bearing the
stigmata and peritremes, is clear evidence that Gretillat has misinterpreted
the dorsal surface of this cosmopolitan parasite of larids.
Genus RALuInyssus Strandtmann
Rallinyssus Strandtmann, 1948, J. Parasit., 34: 512; Wilson, 1965, Pacif.
Insects, 7: 623. Type-species Rallinyssus caudistigmus Strandtmann, 1948,
Loc. cit., 512. Rallinyssoides Fain, 1960, Bull. Annls Soc..r. ent. Belg.,
96: 295. Type-species Rallinyssus congolensis Fain, 1956, Revue Zool. Bot.
afr., 58: 396; 1957, Annls Mus. r. Congo belge Sér. 8vo, 60: 58.
Key to females of Australian species of RALLINYSSUS
il. Gircumanal: striae apsenitiws. are Wie iD © ee ria AGEs ee ohelenc eters ec mec 2
Circuimanaletrille presemtmes notaries vitae sole sitls meds Caner eeu ert. nl eNeb MRA Aer bi5! 0) > 3
ZG) tOpIsthOnotaler shields spkesenity sir sr srism sents ieen anes amaurornmis Wilson
Opisthonotal shield Vabsemtey. seats ao vec she os Slanice e eke e G congolensis Fain
‘3 (1). Podonotal shield about as long as wide, with margin extended laterally beyond
sublateral row of muscle insertions. First pair of setae behind podonotal
shield closely set, not separated by midposterior convexity of shield........
sis ON OE ORE OT Oe RED ONE SO NE RE 6 PO Pe gallinulae Fain
Podonotal shield decidedly longer than wide, with margin not extended laterally
beyond sublateral row of muscle insertions. First pair of setae behind
podonotal shield widely set, separated by midposterior convexity of shield
LM AR Mee al Ra ae abe CAAA aTiasina sobs CU ee Tobe aie rane caudistigmus Strandtmann
-RALLINYSSUS AMAURORNIS Wilson
Rallinyssus amaurornis Wilson, 1965, Pacif. Insects, 7: 624; 1967, Philipp.
J. Sew., 95: 215; 1968, J. med. Ent., Ds 221.
Previous r pee evtatenrered crake, Poliolimnas cinereus (Vieillot)
(Rallidae, Gruiformes).
RALLINYSSUS CONGOLENSIS Fain
Rallinyssus congolensis Fain, 1956, Revue Zool. Bot. afr., 53: 396; 1957,
Annls Mus. r. Congo belge Sér. S8vo, 60:58. Rallinyssus porzanae Wilson, 1967,
Philipp. J. Sci., 95: 215. New synonymy.
Previous record.—Spotless crake, Porzana tabuensis (Gmelin) (Rallidae,
Gruiformes).
In view of the variation of the podonotal shield discussed under the
following species, the two taxa documented above are also synonymized.
RALLINYSSUS GALLINULAE Fain
Rallinyssus gallinulae Fain, 1960, Bull. Annis Soc. r. ent. Belg., 96: 295;
Domrow, 1965, Acarologia, 7: 450; Wilson, 1965, Pacif. Insects, 7: 634; 1968,
J. med, Ent., 5: 221. Rallinyssus rallus Wilson, 1965, Pacif. Insects, 7: 631.
New synonymy.
Previous records (all Rallidae, Gruiformes).—Lewin water-rail, Rallus
pectoralis Temminck. Banded landrail, Hypotaenidia philippensis (Linnaeus),
Innisfail. Eastern swamphen, Porphyrio melanotus Temminck.
Australian specimens show the podonotal shield, and its accompanying
four shieldlets and sixteen setae, arranged as in Wilson’s Fig. 4B. However,
the shield, while longer than wide, is rounded laterally, and therefore inter-
mediate between Wilson’s eroded, and Fain’s subcircular form (Fain’s Fig.
1, where, incidentally, there is also a trace of the second pair of accessory
shieldlets on the right hand side).
324 THE NASAL MITES OF QUEENSLAND BIRDS
Having established this synonymy, it should be pointed out that, by
accepting even further reduction of the podonotal shield as still falling
within the range of intraspecific variation, the question is raised of the
synonymy of the following species (which has priority), and even of RK.
verheyeni Fain (1963d), described from Rallus Linnaeus. There is nothing in
the way of host-specificity to argue against such a step (see Section V below),
and the concept of a single, cosmopolitan species with a circumanal frill,
from a wide range of rallids, is not unattractive.
In view of the peculiarly displaced stigmata in this genus, one might
postulate that the mites assume some definite spatial orientation in the
nasal passages of their hosts.
RALLINYSSUS CAUDISTIGMUS Strandtmann
Rallinyssus caudistigmus Strandtmann, 1948, J. Parasit., 34: 512;
Bregetova, 1951, Parazit. Sb., 13: 118; Domrow, 1966, Proc. Linn. Soc.
N.S.W., 90: 196; Wilson, 1968, J. med. Ent., 5: 221.
Previous records (both Rallidae, Gruiformes).—Dusky moorhen,
Gallinula tenebrosa Gould, Esk. Coot, Fulica atra Linnaeus.
Genus RHINONYsSsUS Trouessart
Rhinonyssus Trouessart, 1894, C. r. Séanc. Soc. Biol., (10) 1: 723. Type-
species Rhinonyssus coniventris Trouessart, 1894, Loc. cit., 724. See also
note below on Sternostomum Trouessart, 1895.
Sommatericola Traigardh, 1904, “Monographie der arktischen Acariden”
(Inaugural Dissertation: Uppsala), p. 28. Type-species Sommatericola
levinseni Trigérdh, 1904, Loc. cit., 29. This genus is based on Somateria
Leach, and Neave (1940) also lists the later spelling Sommateria Kaup.
However, as both these spellings antedate Tragardh’s paper, and it is unlikely
he made a change from Somateria on his own accord, no change in his
spelling seems necessary.
Key to females of Australian species of RHINONYSSUS
alle Anal shield present. Distal segments of palpi forming a compact cone barely
larger than strocha@mter oc. 5 sapovecus Spauae te siete cqeyers depoueis tere a eee oe 4
Anal shield obsolescent. Distal segments of palpi cylindrical, noticeably
longerithan trochanter ~.s.sciscccckcwn ce be els Ons tie aie eee a
2 (1). Podonotal shield entire. Setae on venter of opisthosoma normal ..............
Ree ls Oe oI a ET SE RRC Pinetree ae Ah ore rhinolethrum (Trouessart)
Podonotal shield fragmented. Setae on venter of opisthosoma in ferm of stout
SPLIUCS ye ccre weteetee scans sp eh ce nape rosie Ue Fei ae as ep poliocephalt Fain
3 (1). Sternal shield absent. Genital shield elongate ........ coniventris Trouessart
Sternal shield present. Genital shield as long as broad .................... 4
4 (3). Podonotal shield with posterior margin subrectilinear. Ventral surface of
opisthosoma with 10-12 pairs of setae .......... himantopus Strandtmann
Podonotal shield with distinct posteromedian extension. Ventral surface of
opisthosoma with considerably more, or considerably fewer setae ...... 5
5 (4). Ventral surface of opisthosoma with 20-30 pairs of setae ....................
BE SOC: clo Lice ote Beary oo tena cesta sphenisci Fain and Mortelmans
Ventral surface of opisthosoma with one pair of setae .... minutus (Bregetova)
RHINONYSSUS RHINOLETHRUM (Trouessart)
Sternostomum rhinolethrum Trouessart, 1895, Revue Sci. nat. appl., 42:
393. This is the spelling given in the title and in the formal listing of the
new name. The usage “rinolethrum” a few lines later is an example of
multiple original spelling and should be corrected under Art. 32. van Eyndhoven
‘ Bee eo the labels bear a third spelling. Also Bregetova, 1951, Parazit.
ROBERT DOMROW 325
Rhinonyssus rhinolethrum, Strandtmann, 1951, J. Parasit., 37: 132;
1956, Proc. ent. Soc. Wash., 58: 137; Fain, 1956, Revue Zool. Bot. afr., 53:
149; 1958, Bull. Soc. r. Zool. Anvers, 9: 9; 1960, Revue Zool. Bot. afr., 61:
108; 62: 91; Mitchell, 1960, SWest. Nat., 5: 107; 1968, J. Parasit., 49: 506;
van Hyndhoven, 1964, Zodl. Meded., Leiden, 39: 308; Wilson, 1964, Pacif.
Insects, 6: 383 ; 1968, J. med. Hnt., 5: 222.
Rhinonyssus rhinolethrus Domrow, 1966, Proc. Linn. Soc. N.S.W., 90:
196. Trouessart was at pains to indicate that he considered the correct
original spelling Sternostoma neuter (Arts 30 and 32,), even to the extent
of publishing (1895) the unjustified emendation Sternostomum (Art. 33:
(v. infra). I had therefore considered rhinolethrum adjectival and amended
its termination under Art. 30. However, I now find rhinolethrum is formed
neither exactly from the adjective 6Acpios (olethrius) nor the substantive
oAcpos (Olethrus), and, in the absence of a clear statement by Trouessart
that his specific name be treated adjectivally rather than substantivally,
revert to the original (indeclinable) spelling rhinolethrum.
Rallinyssus rhinolethrum Fain, 1962, Bull. Annls Soc. r. ent. Belg.,
98: 265. As this entry is immediately preceded by the heading “Genre
Rhinonyssus .. .”, the slip of the pen is obvious, and should therefore be
corrected under Art. 32. It should not, presumably, be treated as a new
combination.
Sommatericola levinseni Tragirdh, 1904, “Monographie der arktischen
Acariden” (Inaugural Dissertation: Uppsala), p. 29. Sternostomum levinseni,
Bregetova, 1951, Parazit. Sb., 13:114. Sternostomum levinsi (sic) Vitzthum,
1935, J. Orn., Lpz., 83:569. Rhinonyssus levinseni, van Eyndhoven, 1964,
Zool. Meded., Leiden, 39: 300. Rhinonyssus Dartevellei Fain and Vercammen-
Grandjean, 1953, Revue Zool. Bot. afr., 48:35. The original spelling contra-
venes Art. 28 and should be corrected to dartevellei under Art. 32.
Previous records (all Anatidae, Anseriformes).—Domestic goose, Anser
anser (Linnaeus). Whistling tree-duck, Dendrocygna arcuata (Horsfield)
Kowanyama. Black duck, Anas superciliosa Gmelin. Garganey teal, Anas
querquedula Linnaeus. Mallard (introduced), Anas platyrhynchos Linnaeus.
New host records—Maned_ goose, Chenonetta jubata (Latham)
(Anatidae), Charleville, 25.1.1967, R. D. and J. S. W. (12). Grey teal, Anas
gibberifrons (Miiller), Charters Towers, 10.iii.1966, H. J. L. (1d). Greenshank,
Tringa nebularia (Gunnerus) (Scolopacidae, Charadriiformes), Kowanyama,
11.xii.1967, R. D. and H. A. S. (1 deutonymph).
Like the single specimen recorded by Strandtmann (1956) from a
coot (Fulica Linnaeus, Rallidae, Gruiformes) in Thailand, the new specimen
from Tringa Linnaeus is also best regarded as a straggler —its host was
taken at a small swamp where ducks abound. Strandtmann’s paper (1958)
on the correlation between gregariousness and _ host-specificity in the
Rhinonyssinae should also be consulted. Hirst (1921a@) compares this species
with R. scoticus, but the latter is a nomen nudum, possibly a discarded
manuscript name for one of the other species of Rhinonyssus described in
the same paper.
RHINONYSSUS POLIOCEPHALI Fain
Rhinonyssus poliocephali Fain, 1956, Revue Zool. Bot. afr., 53: 149;
1957, Annls Mus. r. Congo belge Sér. 8vo, 60: 45; Domrow, 1965, Acarologia,
7: 450.
Previous record.—Little grebe, Podiceps ruficollis (Vroeg) (Podicipidae,
Podicipiformes), Esk. Also Kowanyama.
326° THE NASAL MITES OF QUEENSLAND BIRDS
RHINONYSSUS HIMANTOPUS Strandtmann
Rhinonyssus himantopus Strandtmann, 1951, J. Parasit., 37: 1386; 1956,
Proc. ent. Soc. Wash., 58: 182; 1959, J. Kans. ent. Soc., 32: 134; Fain,
1956, Revue Zool. Bot. afr., 58: 149; 1957, Annls Mus. r. Congo belge Sér. 8vo,
60:44; 1958, Bull. Soc. r. Zool. Anvers, 9:10; Haplor. Parc natin. Albert
deux. Sér., 6: 9; 1960, Revue Zool. Bot. afr., 62: 92; 1964, Annls Mus. r. Afr.
cent. Sér. 8vo, 132: 137; Gretillat, 1961, Vie Milieu, 12: 155; Domrow, 1966,
Proc. Linn. Soc. N.S.W., 90: 195. Rhinonyssus strandtmanm Fain and
Johnston, 1966, Bull Soc. r. Zool. Anvers, 38: 25. New synonymy.
Previous records (all Charadriidae, Charadriiformes).— Red-kneed
dotterel, Hrythrogonys cinctus Gould, Kowanyama. Spur-winged plover,
Lobibyx novaehollandiae (Stephens), Esk. Masked plover, ZL. miles
(Boddaert), Kowanyama. Black-fronted dotterel, Charadrius melanops
Vieillot, Kowanyama.
New host record.—White-headed stilt, Himantopus leucocephalus Gould
(Recurvirostridae, Charadriiformes), Kowanyama, 27.x.1966, R. D. and
Ra We (2 29>)!
Strandtmann (1951, 1959) has figured two forms of this species. The
original series from H. meaxicanus (Miller) shows the margin of the
podonotal shield behind the level of the stigmata almost rounded, but with
indications of three shallowly convex lobes. The second form, from Charadrius
vociferus Linnaeus, shows the same bell-shaped shield, but with the postero-
lateral corners angulate and the medial curve less pronounced.
Fain and Johnston (1966) also figured a specimen from (. vociferus,
but show the posterolateral margin somewhat eroded.
The present series from H. lewcocephalus shows the rounded, and the
specimens recorded by Domrow (1966b) from Lobibyx miles and UL.
novaehollandiae the angulate form. However, I can now refer the specimens
from Hrythrogonys cinctus and C. melanops to the eroded, and not to the
original rounded form merely because they were non-angulate (Domrow,
19666). I therefore accept all these specimens as falling within the range
of individual variation of a single species.
RHINONYSSUS SPHENISCI Fain and Mortelmans
Rhinonyssus sphenisci Fain and Mortelmans, 1959, Bull. Soc. r. Zool.
Anvers, 12: 18. Rhinonyssus sphenisci schelli Fain and Hyland, 1963, Bull.
Soc. r. Zool. Anvers, 32: 4. New synonymy. Rhinonyssus schelli, Wilson,
1967, Antarctic Res. Ser., 10: 41.
Previous record.—Adélie penguin (vagrant), Pygoscelis adeliae (Hombron
and Jacquinot) (Spheniscidae, Sphenisciformes).
The synonymy is obvious from the descriptions, and both records are
RHINONYSSUS MINUTUS (Bregetova)
Sternostomum minutus Bregetova, 1950, Dokl. Akad Nauk SSSR, 71:
1007. Sternostoma minutus, Furman, 1957, Hilgardia, 26: 483. Rhinonyssus
minutus, Fain, 1961, Acarologia, 3: 514; Domrow, 1965, Acarologia, 7: 450.
Rhinonyssus pluvialis Fain and Johnston, 1966, Bull. Soc. r. Zool. Anvers,
38: 27. New synonymy.
Previous records (all Charadriidae, Charadriiformes).—Eastern golden
plover, Pluvialis dominicus (Miller). Red-capped dotterel, Charadrius
alexandrinus Linnaeus, Half Tide. Also Tin Can Bay and estuary of Topsy
Ck. Ringed plover (vagrant), C. hiaticula Linnaeus.
The synonymy is obvious from the descriptions, and all hosts are
charadriiforms.
ROBERT DOMROW
4
RHINONYSSUS CONIVENTRIS Trouessart
(Figs 48-57)
‘Rhinonyssus coniventris Trouessart, 1894, C. r. Séanc. Soc. Biol., (10)
1; 724; Hirst, 1921, Proc. zool. Soc. Lond., 1921: 361; Strandtmann, 1951,
J. Parasit., 37: 180; 1956, Proc. ent. Soc. Wash., 58: 180; Mitchell, 1961,
SWest. Nat., 6 108; Ea 1963, Bull. Annls Soc. r. ent. Belg., 99: 88; Domrow,
1965, euiee T: 45( Rhinonyssus echimipes Hirst, 1921, Proc. zool. Soc.
Lond., 1921: 359. eae eae neglectus Hirst, 1921, Proc. zool. Soc. Lond.,
1921: 359; Bregetova, 1951, Parazit. Sb., 13: 117. Rhinonyssus tringae Fain,
19638, Bull. Annls Soc. r. ent. Belg., 99: 96. Rhinonyssus sp. Domrow, 1967,
Proc. Linn. Soc. N.S.W., 91: 216.
Previous records (all Charadriiformes).—Turnstone, Arenaria interpres
(Linnaeus) (Charadriidae). Grey plover, Squatarola squatarola (Linnaeus)
(Charadriidae). Eastern golden plover, Pluvialis dominicus (Miiller) (Char-
adriidae). Red-capped dotterel, Charadrius alexandrinus Linnaeus, Half
Tide. Also Tin Can Bay. Ringed plover (vagrant), Charadrius hiaticula
Linnaeus. Wood-sandpiper, 7'ringa glareola Linnaeus (Scolopacidae). Sander-
ling, Crocethia alba (Vroeg) Scolopacidae). Dunlin (vagrant), Hrolia alpina
(Linnaeus) (Scolopacidae). Knot, Calidris canutus (Linnaeus) (Scolo-
pacidae).
New host record.—Red-necked stint, Hrolia ruficollis (Pallas) (Scolo-
pacidae), estuary of Topsy Ck., 29.x.1966, R. D. (12).
Two considerations underline the artificiality of distinguishing, even
as subspecies, the three nominal taxa of Trouessart and Hirst, whose hosts
include a range of charadriiform birds. On a morphological level, Strandtmann
(1951), studying material from Trouessart’s type-host, A. interpres, figured
a female with a coniventris-like, and a male with a neglectus-like podonotal
shield. Indeed, the degree of erosion of the podonotal shield shows clinal
variation in the direction R. echinipes—R. coniventris—R. neglectus, but a
convincingly identical pattern of muscle insertions is retained in all three.
Further, Fain (1963@), in an addendum, records specimens from the same
host, which bridge another morphological gap (setation of femur IV) in
the first couplet of his key attempting to separate the three forms.
On an ecological level, Fain (1968a@) lists R. c. echinipes from Charadrius
alexandrinus in Europe, and suggests this form is host-specific for Charadrius
Linnaeus. However, in Queensland, two further series (89 2 and 1é, Tin
Can Bay, 29.vi and 17.viii.1966, R. D. and J. S. W.) confirm my earlier
record (1965c) of a neglectus-like form from C. alexandrinus.
The female from FH. ruficollis recalls R. tringae in the shape of its
podonotal shield, but perhaps even here the same pattern of muscle insertions
noted above can be made out (Fig. 56), and I therefore maintain the
synonymy. (A paratype of R. tringae agrees well with Fain’s illustration,
but shows a distinct setal remnant voWe Es the rear of each arm of the
podonotal shield.)
The new Queensland specimen is iMemeated in detail (Figs 48-57)
because its setation (e.g. on the sternal shield and tarsus IV) differs
somewhat from that of Fain’s material. Apart from tarsus I, which is
identical ventrally with Fain’s Fig. 10, leg I is similar to leg II, except
that (i) only two filamentous setae are present on the trochanter; (it)
seven minute setae are present ventrally on the femur; and (#7) two additional
minute setae are present on the ventrodistal margin of the genu. The setal
formulae, therefore, are: coxae 2.2.2.1; trochanters 4.5.5.5; femora 11.10/9.5.7;
genua 10.8.10.9; tibiae 9.9.8.9; tarsi-.18.18.18 (mv present).
328. THE NASAL MITES OF QUEENSLAND BIRDS
A species such as R. coniventris, morphologically much reduced and with
a world-wide distribution throughout a group of birds as extensive as the
Charadrii, may be expected to show a wider range of individual variation
than a more zoogeographically restricted species.
Figs 48-55. Rhinonyssus coniventris Trouessart (Q from Erolia ruficollis) —48-49,
Ventral and dorsal views of idiosoma. 50-55, Ventral and dorsal views of legs II-IV.
ROBDRT DOMROW 329
Genus RuANDANYsSsUS Fain
Ruandanyssus Fain, 1957, Annls Parasit. hum. comp., 32: 148. Type-
species Ruandanyssus terpsiphonet Fain, 1957, Loe. cit., 148.
Key to females of Australian species of RUANDANYSSUS
il Opisthonotal shield considerably narrower than podonotal shield, and tapering
posteriorly. Most idiosomal setae reaching at least half way to the base
of the next nearest seta. Tritosternum with fully formed laciniae........
RUETaneT Cuotharam ie eiver citewencre ferme chore ber at scegs ei niaves aero Ycuelprrtersccvansheronotere aha aie. 6 lterpsiphonei Fain
Opisthonotal shield almost as wide as podonotal shield, and somewhat truncate
posteriorly. Most idiosomal setae falling far short of the base of the
next nearest seta. Tritosternal laciniae obsolescent ........ artami, D. sp
RUANDANYSSUS TERPSIPHONEI Fain
(Figs 58-68)
Ruandanyssus terpsiphonei Fain, 1957, Annls Parasit. hum. comp., 32:
148; 1960, Revue Zool. Bot: afr., 62: 98; Domrow, 1965, Acarologia, 7: 432.
Ruandanyssus terpsiphonei terpsiphonei, Sakakibara, 1968, J. med. Ent.,
5: 17. Ruandanyssus terpsiphonei echongi Sakakibara, 1968, J. med. Ent.,
5:15. New synonymy.
Previous records (all Passeriformes).—Spectacled flycatcher, Monarcha
trivirgata (Temminck) (Muscicapidae), Mt. Jukes. Also Innisfail. Black-
faced cuckoo-shrike, Coracina novaehollandiae (Gmelin) (Campophagidae),
Ksk. Little cuckoo-shrike, C. robusta (Latham), Esk. Rufous whistler,
Pachycephala rufiventris (Latham) (Pachycephalidae), Esk and Mt. Jukes.
Apostle-bird, Struthidea cinerea Gould (Corvidae), Condamine.
New host records (all Passeriformes).—Leaden flycatcher, Myiagra
rubecula (Latham) (Muscicapidae), Brisbane, 15.ix.1965, Esk, 13.11.1968, and
Innisfail, 3.vili.1965, R. D., B. H. K., and J. S. W. (162 2, 44 ¢, and 1
protonymph). White-winged triller, Lalage tricolor (Swainson) (Campo-
phagidae), Kowanyama, 25.x.1966, R. D. (22 2,16). White-browed scrub-
wren, Sericornis frontalis (Vigors and Horsfield) (Sylviidae), Esk, 6.x.1966,
R. D. and J. 8S. W. (182 29,16). Masked wood-swallow, Artamus personatus
(Gould) (Artamidae), Charleville, 24.1.1967, R. D. and J. S. W. (12 ). Golden
whistler, Pachycephala pectoralis (Latham) (Pachycephalidae), Esk,
14.vii.1965, R. D. and J. S. W. (152 2,46 ¢,1 protonymph). Grey whistler,
P. griseiceps Gray, Innisfail, 8.vii, and 1 and 10.ix.1965, G. J. B. and H. I. McD.
(862 2,56 6,1 proto-, and 1 deutonymph).
As presaged by the absence of one genital seta, the female I illustrated
(1965c) is confirmed as an extreme individual variant lacking both metasternal
setae. On checking a long series of both sexes, both metasternals were
invariably found present (as originally figured for the male), except in one
Specimen, where one is lacking (Figs 67-68).
Further, an examination of 75 specimens yielded no aberrancies from
the 1.1 discal setae figured by Fain and myself on the opisthonotal shield
except 1.0 three times (Figs 58-60). However, the material from Myiagra
rubecula differs in showing additional longitudinally arranged pairs of discal
setae on the opisthonotal shield. The following formulae were noted: 1.8
once, 2.2 once, 2.3 four times, 3.3 nine times, 3.4 three times, and 4.4 once
_ (Figs 61-66). I do not feel these minor varieties merit a name.
D
330 THE NASAL MITES OF QUEENSLAND BIRDS
RUANDANYSSUS ARTAMI, N. Sp.
(Figs 69-74, 161-164)
Ruandanyssus sp. Domrow, 1967, Proc. Linn. Soc. N.S.W., 91: 217.
Diagnosis.—k. artanvi is readily separable from the preceding, and only
other known species, R. terpsiphonei Fain (1957a), by using the key above.
as
Ny
Q)
a 68 | :
Seri ok ye el of
Figs 56-57. Rhinonyssus coniventris Trouessart (2 from Hrolia ruficollis) —56, Podonotal
shield and mesonotal shieldlets. 57, Ventral view of capitulum (right palp in dorsal
view ).
Figs 58-68. Ruandanyssus terpsiphonei Fain.—58, Opisthonotal shield of ¢ from Monarcha
trivirgata. 59-60, Opisthcnotal shield of 92 from Pachycephala rufiventris and Sericornis
frontalis. 61-62, Opisthonotal shield of ¢¢ from Myiagra rubecula. 63-66, Opisthonotal
shield of 99 from M. rubecula. 67-68, Sternal and genital shields of 99 from P. griseiceps.
ROBERT DOMROW 331
Types.—Holotype female, allotype male, and two paratype females from
black-faced wood-swallows, Artamus cinereus Gould (Artamidae, Passeri-
formes), Kowanyama, 27.iii and 1.iv.1965, R. D. and J. 8. W. Holotype
and allotype N. I. C.; paratypes R. D and A. F.
Also 82 2,124, and 2 protonymphs from a little wood-swallow, A. minor
Vieillot, Charleville, 23.1.1967, R..D. and J.S. W.
Female.—Idiosoma 585-595. long in little compressed specimens (one
containing embryonic larva, another with undeveloped ovum extruded),
Figs 69-74. Ruandanyssus artami, n. sp. (from Artamus cinereus) —69-70, Ventral and
. dorsal views of idiosoma of 9. 71, Chelicera of 2. 72-73, Dorsal and ventral views of
capitulum of 9. 74, Ventral view of idiosoma of J.
aoe THE NASAL MITES OF QUEENSLAND BIRDS
slightly constricted behind stigmata. Podonotal shield (Fig. 70) as long
as wide, rather evenly rounded anteriorly and laterally, but with posterior
margin weakly trilobate; with nine pairs of setae (only midposterior pair
prominent), and two pairs of submarginal pores. Opisthonotal shield concave
anteriorly, roundly convex posteriorly, and trilobate laterally; with one
pair of small setae discally in addition to terminal pygidial pair. Both
shields closely granulate (except for narrow marginal strip), and further
marked by paired muscle insertions. Dorsal cuticle with nine pairs of
setae of varying size. Stigmata provided with short peritremes and followed
by lateroventrally-directed poststigmatic shields.
Sternal shield (Fig. 69) subtriangular, well sclerotized, granulate, but
with indefinite margins; bearing SI (with attendant pores) and flanked
by SII-IIT; metasternal setae present, but small. Genital shield drop-shaped,
weakly granulate, with some longitudinal strengthening discally, and rounded,
rayed operculum; genital setae small. Anal shield longer than wide, well
rounded at front, and denser at sides; cribrum present. Anus set well
forward, with adanal setae near its anterior edge; postanal seta smaller,
set well back from anus. Ventral cuticle with twelve setae arranged 2.4.6.
All body setae simple, with rather blunt tips.
Leg setae similar (Figs 163-164). Coxae 2.2.2.1. Trochanter I (1—0/2-1),
II (1-1/2-1), II-IV (1-0/2-1). Femur I (2-3/1-2), IT (1-4/2-1), Ill
(1-4/1-0), IV (1-4/0-0). Genu I (1-4/1-1), II-III (1-4/0-1), IV (1-3/0-0).
Tibia I (1-4/2-1), II-IV (1-8/2-1). Tarsi —.17.17.16 (mv lacking on II-IV;
ad2 also lacking on IV). Minor variants common on femora (I with 1 pl,
IL with 1 v or 0 pl, IIT with 0 v, IV with 8 d), and tarsi (III lacking al2 or
ave, IV lacking alz or al3). Tarsus I with 21 setae in addition to dorsodistal
sensory islet of five rods (Figs 161-162). Two lyriform fissures present,
one dorsally and one posterolaterally (1) or anterolaterally (II-IV), between
basi- and telofemora. Tarsi II-IV with four lyriform fissures, one dorso-
basally on basitarsus, one middorsally on telotarsus, and one dorsally
(divided) and one ventrally between basi- and telotarsus. Tarsus I with
two fissures only (first and last of above four). Ambulacrum I similar to
IL-LV, but pretarsus slightly stouter.
Basis capituli (Figs 72-73) with indication of deutosternum, but without
denticles; with two capitular setae. All six hypostomal setae present, but
small. Epistome hyaline marginally, with denser dendritic area discally.
Palpi with usual five free segments; setal formula 0.3.4.7. Genu with
dorsobasal lyriform fissure. Tarsus obscured dorsally by tibia; with about
six small setae. Claw bifid. Cheliceral shaft of uniform diameter; single
(movable) digit faleate, occupying one-sixth of total length (Fig. 71).
Tritosternum hyaline, with base well developed, but laciniae obsolescent.
Male.—Idiosoma 485, long in slightly compressed specimen. Essentially
as in female except for fusion of sternal, metasternal, and genital complexes
(Fig. 74), and presence of spermatodactyl on chelicerae.
Protonymph.—Podonotal shield as in female, but with only eight pairs
of setae (verticals lacking). Pygidial shield indistinct, but with two barbulate
setae twice as strong as strongest of simple dorsal setae. Setal arrangement
on dorsal cuticle as in female, but including pair of setae taken in discally
by expansion of pygidial shield into opisthonotal shield. Stigmata with
peritremes, but unarmed.
Sternal shield with two pores near SI; SI-III submarginal. Anal
shield as in female. Ventral cuticle with eight setae arranged 2.4.2.
Chaetotaxy of coxae, genua, tibiae, and tarsi as in adult (minute
ventrodistal setae on plantar surface of tarsus I difficult to see even at x 500),
ROBHRT DOMROW 333
with following individual variation: coxa III lacking pv once, and genu IV
(1-2/0-0) twice. Trochanters as in adult, except [IV (1-0/2-0), with following
individual variations: IT (1-0/2-1) once, and III (1-0/2-0) once. Femur I
as in adult, II (1-4/1-0), III (1-3/1-0), IV (1-8/0-0), with following
individual variation: IT (1-4/1-1) once. Lyriform fissures arranged as in
adult, but dorsobasal fissure on basitarsus [I absent, and dorsal fissure
between basi- and telotarsi II-lV undivided.
Capitulum as in female, but with five deutosternai denticles in single
file.
Genus RuiNorcius Cooreman
Rhinoecius Cooreman, 1946, Bull. Mus. r. Hist. nat. Belg., 22: 1. Type-
species Rhinoecius oti Cooreman, 1946, Loc. cit., 1. Zumptnyssus Fain, 1959,
Bull, Annls Soc. r. ent. Belg., 95: 112. Type-species Ruandanyssus buboensis
Fain, 1958, Revue Zool. Bot. afr., 58: 292. New synonymy.
The presence or absence of a_ tritosternum, as indicated under
Tinaminyssus above, and Ptilonyssus below, has no meaning at a generic level.
Further, both genera are restricted to owls.
Key to females of Australian species of RHINOECIUS
1. Podonotalishieldwentinemenorccs cee cece cee einer cooremani Strandtmann
Podonotal shield fragmentary, with one larger anterior, and four smaller
WOSTEEIONE LU ASM ENTS aris ers ec aac cae at oe ore) a oie ales tytonis Fain
RHINOECIUS COOREMANI Strandtmann
(Figs 75-77)
Rhinoecius cooremani Strandtmann, 1952, Proc. ent. Soc. Wash., 54:
208.
This genus, previously recorded only from strigid owls in Europe, Africa,
and the U.S.A., may now be added to the Australian fauna: 92 2 and 1
protonymph from a boobook owl, Ninox novaeseelandiae (Gmelin) (Strigidae,
-Strigiformes), Esk, 5.11966, R. D. and J.S.W.
I have examined paratypes of Strandtmann’s three species (two of
which have been further illustrated by do Amaral, 1962), and my specimens
differ from his Rk. cooremani only in showing (i) the anteriormost pair of
dorsal cuticular setae apparently set on the margin of the podonotal shield
(Fig. 77) (1 say “apparently” because the cuticle on the specimens available
is somewhat exfoliated, and could have come, together with its setae, to
lie over the edge of the shield); and (i) the postanal seta present, but
rather weaker than the adanals (Figs 76-77). The capitulum is depicted
in Fig. 75.
RHINOECIUS TYTONIS Fain
Rhinoecius tytonis Fain, 1956, Revue Zool. Bot. afr., 53: 394; 1957,
Annls Mus. r. Congo belge Sér. 8vo, 60: 131, 138.
Previous record.—Barn-owl, Tyto alba (Seopoli) (Tytonidae, Strigi-
formes).
Genus Prinonyssus Berlese and Trouessart
Ptilonyssus Berlese and Trouessart, 1889, Bull. Biblioth. scient. Ouest,
2: 128. Type-species Ptilonyssus echinatus Berlese and Trouessart, 1889,
Loc. cit., 129. Rhinonyssoides Hirst, 1921, Proc. zgool. Soc. Lond., 1921: 770.
Type-species Rhinonyssoides trouessarti Hirst, 1921, Loc. cit., 770. Neonyssus
Hirst, 1921, Proc. Zool. Soc. Lond., 1921:771. Type-species Neonyssus
intermedius Hirst, 1921, Loc. cit., 771. Neonyssoides Hirst, 1923, Proc. zool.
304 THE NASAL MITES OF QUEENSLAND BIRDS
Soc. Lond., 1923: 975. Type-species Rhinonyssus (Neonyssoides) nucifragae
Hirst, 1923, Loc. cit., 975. Hirst’s three type-species have since been described
and illustrated by Fain (1960a), Fain and Hyland (19626), and Bregetova
(1967). Ptilonyssoides Vitzthum, 1938), J. Orn., Lpz., 83: 581. Type-species
Ptilonyssoides triscutatus Vitzthum, 1935, Loc. cit., 581. Rhinacarus de Castro,
1948, Archos Inst. biol., S Paulo, 18: 257, nec Nehring, 1884, Sber. Ges.
75
Figs 75-77. Rhinoecius cooremani Strandtmann (@2 from Ninox novaeseelandiae) —75,
Ventral view of capitulum (right palp in dorsal view). 76-77, Ventral and dorsal
views of idiosoma.
Figs 78-80. Ptilonyssus neochmiae, n. sp. (2 from Neochmia phaeton).—78, Ventral
view of capitulum (right palp in dorsal view). 79-80, Dorsal and ventral views of
tarsus III.
Fig. 81. Ptilonyssus cinnyris Zumpt and Till (? from Cyrtostomus frenatus) .-Opistho-
notal shield.
ROBHRT DOMROW 335
naturf. Freunde Berl., 1884: 64. Type-species Rhinonyssus (Rhinacarus)
angrensis de Castro, 1948, Loc. cit., 257. New synonymy. The relevant
paragraph of Nehring runs: “Der Name Halarachne erscheint mir nicht sehr
gliicklich gewdhlt, da diese Milbe nach den bisherigen Beobachtungen
keineswegs frei im Meerwasser lebt, sondern ihre Eaxistenz an die Nasenhéhle
der Kegelrobben (vielleicht auch anderer Pinnipedier) bindet wnd_ sich
vermuthlich nur von Thier zu Thier iibertragt. Ich wiirde sie Rhinixodes oder
Rhinacarus nennen, wenn sie new zu benennen wire.’ Nehring’s use of the
subjunctive indicates that he did not mean to erect these names for possible
use in the future, but that he was only indicating what names he himself
might have used had he been naming the taxon. Indeed, he did not even
show a preference for either name, and it could be argued that the names
were not proposed for taxonomic use (Art. 1). On the other hand, assuming
these names were proposed for taxonomic use, they can only be considered
as synonyms of Halarachne Allman, as did Nehring himself, since he continued
to use Halarachne. Thus, as names first published as synonyms, they are
unavailable under Art. 11.
Flavionyssus de Castro, 1948, Archos Inst. biol., S Paulo, 18: 266. Type-
species Ptilonyssus (Flavionyssus) rabelloi de Castro, 1948, Loc. cit., 266.
Rochanyssus de Castro, 1948, Archos Inst. biol., S Paulo, 18: 272. Type-
species Neonyssus (Rochanyssus) werneri de Castro, 1948, Loc. cit., 272.
Paraneonyssus de Castro, 1948, Archos Inst. biol., S Paulo, 18: 274. Type-
species Neonyssus (Paraneonyssus) enriettii de Castro, 1948, Loc. cit., 274.
Travanyssus de Castro, 1948, Archos Inst. biol., S Paulo, 18: 276. Type-species
Neonyssus (Travanyssus) paranensis de Castro, 1948, Loc. cit., 276. Vitenyssus
de Castro, 1948, Archos Inst. biol., S Paulo, 18: 277. Type-species Dermanyssus
niteschi Giebel, 1871, Z. ges. Naturw., 38: 31. New synonymy. Cas Baker and
Wharton, 1952, “An introduction to acarology” (Macmillan: New York),
p. 81 (unnecessary nomen novum for Rhinacarus de Castro). Type-species as
for Rhinacarus de Castro, v. supra. New synonymy. Astridiella Fain, 1957,
Riv. Parassit., 18: 94. Type-species Ptilonyssus scotornis Fain, 1956, Revue
Zool. Bot. afr., 53: 148. New synonymy. Hapalognatha Butenko, 1959, Nauch.
Dokl. vyssh. Shk., 2:17; 1960, Zool. Zh., 39: 1490. Type-species Hapalognatha
prima Butenko, 1959, Loc. Cit., 17 (nomen nudum under Art. 13); 1960,
Loc. cit., 1490. Passeronyssus Fain, 1960, Revue Zool. Bot. afr., 61: 110;
1962, Tbid., 66: 139; Fain and Nadchatram, 1962, Bull. Annls Soc. r. ent.
Belge., 98: "O75. Type- -species Ptilonyssus viduae Fain, 1956, Revue Zool. Bot.
afr., 53:147; 1957, Annls Mus. r. Conge belge Sér. 8vo, 60: 120. New synonymy.
Tyranninyssus Brooks and Strandtmann, 1960, J. Parasit., 46: 418. Type-
species T'yranninyssus tyrannus Brooks and Strandtmann, 1960, Loc. cit.,
419. New synonymy. Locustelionyssus Bregetova, 1964, “Some problems of
evolution of the rhinonyssid mites” (Nauka: Leningrad). Locustellonyssus
is not mentioned in the text of this article, but occurs in an accompanying
photograph of a table on host-specificity. This does not constitute publication
under Art. 8; and, in any case, as no type-species is given, Locustellonyssus
is unavailable here under Art. 13. Also 1965, Zool. Zh., 44: 1093. Type-species
Locustellonyssus amurensis Bregetova, 1965, Loc. cit., 1093. New synonymy.
Periglischrodes Baker and Delfinado, 1964, Pacif. Insects, 6: 589. Type-species
Periglischrodes gressitti Baker and Delfinado, 1964, Loe. cit., 589. Otocorinyssus
Bregetova, 1967, Parazit. Sb., 28: 127. Type-species Neonyssus (Otocorinyssus)
melanocoryphae Bregetova, 1967, Loc. cit., 127. New synonymy. Frigilonyssus
Bregetova, 1967, Parazit. Sb., 23: 130. Type-species Ptilonyssus coccothraustis
Fain and Bafort, 1963, Bull. Annis Soc. r. ent. Belg., 99: 447. New synonymy.
Spizonyssus Bregetova, 1967, Parazit. Sb., 23: 131. Type-species Ptilonyssus
serinit Fain, 1956, Revue Zool. Bot. afr., 53: 1389; 1957, Annis Mus. r. Congo
336: THE NASAL MITES OF QUEENSLAND BIRDS
belge Sér. 8vo, 60: 90. New synonymy. Neotyranninyssus Fain and Aitken,
1967, Bull. Inst. r. Sci. nat. Belg., 43: 27. Type-species Neotyranninyssus
fluvicolae Fain and Aitken, 1967, Loc. cit., 29. New synonymy. Trochilonyssus
Fain and Aitken, 1967, Bull. Inst. r. Sci. nat. Belg., 43: 29. Type-species
Trochilonyssus trinitatis Fain and Aitken, 1967, Loc. cit., 32. New synonymy.
Pipronyssus Fain and Aitken, 1967, Bull. Inst. r. Sci. nat. Belg., 43: 36.
Type-species Pipronyssus manaci Fain and Aitken, Loc. cit., 36. New
synonymy.
The number of synonyms for Ptilonyssus s. l. is already large, but I make
no apology for increasing it further. The reason that the boundaries between
these taxa have been drawn, and redrawn, is the simple one that they
correspond to nothing at all, their only purpose having been to divide,
in the light of arbitrary and trivial morphological characters, a group of
which all other considerations emphasize the fundamental unity. The smaller
the taxon, the greater should be the morphological and ecological gap
between it and its nearest relatives.
It has proved difficult to arrange this large series in a completely orderly
system, and I have therefore compromised between the order of the key and
the order of the hosts. This point is discussed further in Section V_ below
on host-specificity.
Key to females of Australian species of PTILONYSSUS
1. Stismatay provided twathy penilremess = eaaeicicie ci octet 2
Susmataynon) provided) with) pPeritreMmles. -e era aceite een nee 49
2 (1). Dorsal armature (excluding mesonotal shieldlets) comprising three units:
podonotal, opisthonotal, and pygidial, of which the latter two may, or
may wot, be fuSeds © s cictoisnt vor, byte oc eriueneretoiene dt alee ae Eee 3
Dorsal armature (excluding mesonota] shieldlets) comprising two . units:
podonotal and pygidial, of which the latter may be obsolescent or
albsemt, Ui CEH OU Gah SS RA SD A. OE) One 14
3 (2). Mesonotal sclerotization with two pairs of strong, peg-like setae in tandem.
Bostanale seta’ AOSEM Ew xcs ce Aen watt ye edsi atic rors oe ailuroedi Domrow
Mesonotal complex otherwise. Postanal seta normally present, but absent in
iP PYGMACUS *(BIreZetOva). “soo. Sea ass ee ed ole eee 4
44¢3).. “‘Cribrum! present? \ 01% ASS red, Bae eee de ek 5
Cribrumisabsent yaaa a. 3 ee Oo nn EERE CR 3 Gio oc 13
5 (4). Pygidial shield fused with opisthonotal shield .................. bce spaueee a, Uae 6
By Sida: ishvel detree. 262i. a che cg Meee inewas ehalte saskatoon ee 12
6 (5). Metasternal setae normally present, but absent in specimens of P. carduelis
Fain from Chloris chloris (Linnaeus)
Metasternal ‘setaelabsent Skeet a7.. tle EDR TRE Ree See 11
7 (6). Podonotal shield with four setae considerably stronger than remainder on
COTS UIs brs, Seats ete: oS oe tecoR ees eae ee eae eae eer carduelis Fain
ANS Setae on dorsum unit orm sine sizer see ee eee ae eae 8
SC), “Podonotal ‘shield “strongly ‘convex laterally ~..-)25-.-0) 02) eee 9
Podonotalgshicld@paralilel-sided) =sese hao ee eee eee malurt Domrow
9 (8). Opisthonotal shield widest anteriorly. Adanal setae near front of anus. Coxal
setae. uniform. mm, lemethy | eS) og 05S ces Steen a oe ee ee 10
Opisthonotal shield widest at its middle. Adanal setae near back of anus.
Sctasonecoxay MVaeclon salem eee an see hirsti (de Castro and Pereira)
HOMEO A MBostanall setay presents AO. +. ok Na ee A Se emberizae Fain
Postanalesetacabsenta aes sees cnn eee pygmaeus (Bregetova), n. comb.
11 (6). Opisthonotal shield subtriangular, with four pairs of setae in addition to
pygidials. Tarsi II-IV with al,, av,, pv,, al,, av., and pv. in form of strong,
slightly curved spurs, which, at their apex, curve distinctly while tapering
Mapidiliye 4! eh AGE. AIAN eae ho Reena ore eeu ce tensor colluricinclae Domrow
Opisthonotal shield subrectangular, with one pair of setae in addition to
pygidials. Tarsi II-IV with al,, av,, pv,, and pl, irregularly inflated....
ES TT RIT OTD en Cece ce Mn tics One CD rae chee capitatus (Strandtmann)
12 (5). Opisthonotal shield fully as wide as podontal, and considerably larger
than pygidial shield. Anus occupying entire anterior angle of elongate
anal shield, preceding all three anal setae ........ triscutatus (Vitzthum)
Opisthonotal shield much narrower than podonotal, and subequal to-pygidial
shield. Anus centrally placed on anal shield, flanked laterally by adanal
setae (tid. Ree lees ae Ope DRC et Oe ee le sittae Fain
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
(4).
(2).
(14).
(15).
(16).
(17).
(18).
(17).
(20).
(21).
(22).
(23).
(24).
(25).
(26).
(27).
(22).
(21).
(20).
ROBERT DOMROW 337
Opisthonotal shield entire, with pygidial complex obliterated ..............
Frags tacos CIAO Ciacci a RRC aca ACUCe RAI S.CAChCunCani NCAA Eat aria struthideae, n. sp.
Opisthonotal shield fragmented, with pygidial remnants possibly present....
fk cash Olea Coibace curt Le ihaG ON ERI RCRERORE CORR.) “SEVCROICE RNC NRO ie fiche corcoracis, N. sp.
Distinct tritosternum with well developed laciniae present .. nitzschi (Giebel)
Tritosternum rarely present, and then represented only by merest remnant,
WHO UTM CHINA CUM rrr Minchin vari a MMnue rs clatgaine’ ele, cictsl via’ eaeelecelpumerasteioire ts 15
Postanal seta normally present, but absent in specimens of P. terpsiphonei
Fain from Carterornis Mathews and Monarcha melanopsis (Vieillot) .. 16
POStHnal ERO tamaDSOl Umer ten ty wien rv Wal csUt ne esgvecaie = lokecits a) sha aausas ales‘ s\0's's 41
Podonotal shield not flanked on each side by small accessory shield
Behe ASsict RADY WOE calacide aL od ts cs CR GNOL DIN Ee CaP ta: a A Meee Meals adawhah ante ai sats ls eleretous 4 8,283. 17
Podonotal shield flanked on each side by small accessory shield ........ 38
Podonotal shield at least as wide as long, with lateral margins concave in
their posterior half, and therefore divergent behind the peritrematalia.
Posterolateral angles of shield full and rounded, resulting in a wide,
Shrallowilivar Obed. pOSLETLOME eat Sune weieuees cat ebenetel aia aiaveus cleus taicneestaletale oie ee 18
Podonotal shield normally clearly longer than wide, but if not, not formed
BSs ADOME, 5 aye scayehare ce Fae oss ROTORS Ac pRTCR wos) Meh aeans Cs ata Daal, dat Watet MEN outs. > os 20
Podonotal shield not sufficiently reduced posteriorly to leave midposterior
pair of setae free in cuticle immediately in front of mesonotal shieldlets
Ne ee COREA OO PE Or DONC OOO oi Doon rome ee o cracticti Domrow
Podonotal shield with posterior margin sufficiently eroded medially to leave
midposterior pair of setae free in cuticle .....................-00-. 19
Pygidial shield present, either entire or divided ............ motacillae Fain
Pycidialmshield absentee. 6 saya aerelotts ieee langei (Butenko), n. comb.
Podonotal shield not sufficiently reduced posteriorly to leave midposterior
_ pair of setae free in cuticle immediately in front of mesonotal shieldlets
BE MTesfe ey Siar ol OE te i ectaN Se Ghose eS Pay TS NS GeTOISI CS SRL HOR, AEROS. CGA 21
Podonotal shield truncate posteriorly, leaving midposterior pair of setae free
in cuticle immediately in front of mesonotal shieldlets .............. 31
Two setae on posterior margin of podonotal shield subequal to remainder on
FSU BUY YE IGE ceteris hE oly ORE DEES aS Te RTS aren gehen oe Sat a 22
Two setae on posterior margin of podonotal shield considerably stronger
thangremamdersony shicldum-anvan ats ae ae ee one es eee 30
Pygidial shield normally entire, but divided in specimens of P. philemoni
from Meliphaga notata (Gould) and M. gracilis (Gould) ............ 23
by eidraleshield wdimidedsorsabsent tise asus eee eee ee ee Ee ee 29
Palpal setae not as long as their corresponding segment. Ventral surface of
OpISthOSomatwithiG-l4epairsvof isctacwm hee. erento eect. eee 24
Four setae on ventrolateral aspect of each palp considerably longer than
their corresponding segments. Ventral surface of opisthosoma with about
ZSEPAINS OL SETAC AL, FAW EIA POOLS OL MR ot oh, balimoensis Sakakibara
All coxal setae normally sharply tipped, but blunt and peg-like on coxa
MOMS EMLCTDSTDNONEis Haine soe, tee eee eee ee eS 25
All coxa] setae inflated, subglobular .................... trouessarti (Hirst)
Genitalisetaeyonwyexenitall ishielde saves meee beets aes cies ster orp, «loa eiste: 26
Genital setae in cuticle adjacent to genital shield ........ philemoni Domrow
All leg setae sharply pointed. Seta mv absent on tarsi II-IV ............ 27
Legs with a mélange of pointed and blunt, peg-like setae. Seta mv present
OU GCA S ee LIV eer eee tenses hice nh sek una Paty ho terpsiphonei Fain
Adanalwsetaceinistronbrotsamisiey artes osc a se eo alco yeuc. do microecae Domrow
Adanalers ctaecumpeltnadeamnSaur a ccc teen catsc eis steren eae seIPre cae aeh-utjcgasreiees aie oa. 28
Podonotal shield larger in posterior half, with posterior margin subrectilinear
and bearing four setae. Metasternal setae present. Ventral surface of
opisthosoma with about six pairs of setae .. nudus Berlese and Trouessart
Podonotal shield larger in anterior half, with posterior margin fully curved
and bearing two setae. Metasternal setae absent. Ventral surface of
opisthosoma with 8-10 pairs of setae ................ orthonychus, D. sp.
Podonotal shield with seven pairs of setae, of which only four are submarginal.
Pygidial shields distinct. Adanal setae in front of anus ..............
ee PTA ER AA RN Ne Log Shae ee sot myzanthae Domrow
Podonotal shield with six pairs of setae, all submarginal. Pygidial shield
obsolescent. Adanal setae behind anus ............ myzomelae Domrow
Adanal setae in front of anus. Pygidial shield entire ...... acrocephali Fain
Adanal setae behind anus. Pygidial shield divided .......... pittae Domrow
TEAC, Same! GINRTRE ooobechsodadaguoteococgsadborrconboprmGuceb co oddopes 32
Tevexitiall Sini@lel GkiwmGle@! OF BUSSNE acecaccouncueocnabondsdcoce sccuscgar 36
338
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
Bt
52
(31).
(32).
(33).
(32).
(31).
(36).
(16).
(38).
(39).
(15).
(41).
(42).
(43).
(44).
(43).
(42).
(47).
(DY):
(49).
(50).
(51).
THE NASAL MITES OF QUEENSLAND BIRDS
Podonotalgshieldvat leastwasylon2egas! wid Gls aceon eine 33
Podonotal shields noticeably. wider chen lone ier ernest rele 35
Podonotal shield convex laterally, with at least six pairs of setae (including
Vertical pain) eAdanale setae tronic Ole aus meee ie eres 34
Podonotal shield concave laterally, with only three pairs of setae (including
vertical pair). Adanal setae behind anus .......... stomioperae Domrow
Podonotal shield with two pairs of setae posteriorly ........ setosaé, 1. sp.
Podonotal shield with one pair of setae posteriorly ...... gerygonae, n. sp.
Setae between podonotal shield and peritrematalia undistinguished. Coxa II
VALE NOW “LVNUETOCMOMSENL [NROGAIS sodbabcoccosoucaundvodonddon dicaei Domrow
Two setae between podonotal shield and peritrematalia on each side consider-
ably stronger than remainder. Coxa II with anterodorsal process ......
CITED EEA SRCICRG Acer on ncee CeIn eaeNIE ere icepAUne eather oN RONG ocer gions haba did oho ruandae Fain
Podonotal shield widest near its middle. Pygidial shields present, even
LED (SIMA ee Oh Sa ie, Bak Ee erty omar «cas Stake. 7) ear a re 37
Podonotal shield clearly widest anteriorly. Pygidial shields completely absent
eae Ce er Cae 3 oreo aE MD ROTI ror Ne chet Caner erence cinnyris Zumpt and Till
Podonotal shield with three pairs of setae in midline (including verticals).
Pygidial shields subcircular, well formed ............ rhipidurae Domrow
Podonotal shield with four pairs of setae in midline. Pygidial shields
CVATIESCEME Se ee. aA ee eel AE Pai en chat ere lymozemae Domrow
Coxal setae inflated, subspherical .......... echinatus Berlese and Trouessart
Coxal ‘setae ‘elongate, slenderly taperine@ <:52::.--.05--+-- 20) oso ee 39
Adanal setalebehind anus’ s.8 Pane See Se, (ae e Rae, See eee 40
Adanallisctachinetront Ob anspor cee om es more cee thymanzae Domrow
Podonotal shield distinctly broader in anterior half, with six pairs of setae
LEENS F445. b, sRodatMtats xnapedanaw.nansramamen skebeuananatichanar ot onatisvaratcclevenetatotete ameter gliciphilae Domrow
Podonotal shield narrowly subrectangular, with four pairs of setae (including
VELEUCAal HPAairy iy |. ein hs ae reat ce rol iaedc Shale dene ere aaa meliphagae Domrow
Py Lidial LSHield) <GMET ES) ce 5 sussws awe ousorsar ostowrerepiok sr ses esol 0) 05 Gon stheyo-8 Waske V2 GH ER RRR SECC aone 42
Pycidials shieldivabsent’ Sct Pal, esate Reese osteo tae ke othe ete) ae cerchneis Fain
Podonotal shield* not sufficiently reduced posteriorly to leave midposterior
pair of setae free in cuticle immediately in front of mesonotal shieldlets
Se ae ee ah RRR POA ANN PE Ay NRRL ME OE Ad bdo Oo oOo ese 6 43
Podonotal shield truncate posteriorly, leaving midposterior pair of setae free
in cuticle immediately in front of mesonotal shieldlets ............ 47
Setae on coxae and venter of opisthosoma slenderly tapering .......... 44
Setae on coxae and venter of opisthosoma truncate, rod-like .......... 46
Coxay il without anterodorsal processignes a4. See Oboe ee eee eee 45
Coxayliawithwanterodorsalaprocess) eee ir eer ren ae eee monarchae, n. sp.
A stout species when engorged. Podonotal shield strongly cordate, with regular
MAT PANS) A ages oye Beso a exon srs ORI TS a Oar sphecotheris Domrow
An elongate species when engorged. Podonotal shield with lateral margins
very irregular, and noticeably narrower in posterior third ..............
a eh SES Senos TT Rca OE MRSS co CEO Ce I OR ee sturnopastoris Fain
Chelicerae attenuate only at extreme tip ............ novaeguineae (Hirst)
Cheliceraeyattenuateminedi Stale hiallissees eee eee een grallinae Domrow
Chelicerae attenuate only at extreme tip ............ novaeguineae (Hirst)
Chelicerae attenuatesinadistalp half. s.4 566 o-oo ae oe ae eee oe 48
Podonotal shield with six pairs of setae. Setae on dorsum of opisthosoma
ina) TOM OF INGATAT SIOUNES sge.cdbcccedoocsdccadasecace psophodae Domrow
Podonotal shield with five pairs of setae. Setae on dorsum of opisthosoma
Wa THOT Oil WMT TROGI Scoscodcoccccctaccesscoosscese: macclurei Fain
Extensive opisthonotal or mesonotal shield present, irrespective of pygidial
COMPO, Oe eicco oye. ee ot MERE RO AERO LOE GLE OCR Eee 50
Only pygidial shield present on opisthonotum .... bradypteri (Fain), n. comb.
Pygidial complex, if present, fused to opisthonotal shield .............. syil
Pyeidial’ “shield” "Giscrete eae sven oc ne ats ee A chee Eee 53
Opisthonotal shield extended back to include pygidial complex. Sternal shield
Waders thianyul ONS yess cea ast ee hatt Et nee ERPS eE Bercaeee neochmiae, N. Sp.
Mesonotal shield not extending back to pygidium; pygidial shield absent.
Syiemnel Shwielel lOmeer Whar WG sosoccsscaccacccsoeccsucccososeocenc 52
Genital setae off shield. Sternal, genital, and ventral setae in form of weak
TOO Sree eas costal ee ee eee elbeli (Strandtmann), n. comb.
Genital setae on shield. Sternal, genital, and ventral setae in form of heavy
DOSSF arte tienen crare ane et nek oe gta ee ee angrensis (de Castro), n. comb.
couplet.
* The nature of the podonotal shield is unknown in P. novaeguineae (Hirst) (see
Fain and Hyland, 1962b), and it is therefore keyed out through both halves of this
ROBERT DOMROW 339
58 (60). Opisthonotal shield elongate, bearing three pairs of setae. SI on shield.
Anal shield evenly sclerotized. Leg setae in form of small spines ........
Pap HO Oey eet CteD coche eh ie 2 EE DRELEACRORCUC PEMD Okcne” «RCRD cr Punt ROPER fac dioptrornis Fain
Opisthonotal shield quadrate, bearing two pairs of setae. SI off shield. Anal
shield with heavily sclerotized band between anus and cribrum. Many
LOLNSCLHO MATEO AMGe NTALCIY 1c) axe cp eaters 010 foieis aus caaere tars flames ee dicruri Fain
PTILONYSSUS MALURI Domrow
Ptilonyssus maluri Domrow, 1965, Acarologia, 7: 451. Neonyssus maluri,
Bregetova, 1967, Parazit. Sb., 23: 138.
' Previous record.—Red-backed wren, Malurus melanocephalus (Latham)
(Sylviidae, Passeriformes), Esk.
PTILONYSSUS COLLURICINCLAR Domrow
Ptilonyssus -colluricinclae Domrow, 1964, Acarologia, 6: 596. Neonyssus
colluricinclae, Bregetova, 1967, Parazit. Sb., 23: 138.
Previous record.—Grey shrike-thrush, Colluricincla harmonica (Latham)
-(Pachycephalidae, Passeriformes), Brisbane. Also Wilson’s Peak and Esk.
New host records.—Rufous shrike-thrush, C. megarhyncha (Quoy and
Gaimard), Innisfail, vi-ix. 1965, G. J. B., R. D., H. I. McD., and J. S. W.
(522 2, 10 protonymphs); Esk, 14.vii.1965, R. D. and J. S. W. (472 9,
5 nymphs). Golden whistler, Pachycephala pectoralis (Latham), Innisfail,
1.ix.1965, H. I. McD. (112 2). Grey whistler, P. griseiceps Gray, Innisfail,
9.vii.1965, G. J. B.and H. I. McD. (192 ).
PTILONYSSUS CAPITATUS (Strandtmann)
(Figs 82-89)
Paraneonyssus capitatus Strandtmann, 1956, J. Kans. ent. Soc., 29: 133.
Astridiella capitatus, Fain, 1959, J. ent. Soc. sth. Afr., 22:21. Neonyssus
(Otocorinyssus) capitatus, Bregetova, 1967, Parazit. Sb., 23: 127.
This species may now be recorded from Australia: 22 2 from a
Horsfield bushlark, Mirafra javanica Horsfield (Alaudidae, Passeriformes),
Kowanyama, -3.iv.1965, R. D. The only previous record is from an American
lark (Otocoris Bonaparte).
PTILONYSSUS CARDUELIS Fain
Ptilonyssus carduelis carduelis Fain, 1962, Bull. Annls Soc. r. ent. Belg.,
98: 253. Neonyssus (Frigilonyssus) carduelis carduelis, Bregetova, 1967,
Paragit. Sb., 23: 130. Ptilonyssus carduelis chloris Fain, 1962, Bull. Annis. Soc.
r. ent. Belg., 98: 257. New synonymy. Neonyssus (Frigilonyssus) carduelis
chloris, Bregetova, 1967, Parazit. Sb., 23: 130.
Previous record.—Greenfinch (introduced), Chloris chloris (Linnaeus)
(Fringillidae, Passeriformes).
PTILONYSSUS EMBERIZAE Fain
Ptilonyssus emberizae Fain, 1956, Revue Zool. Bot. afr., 53: 140; 1957,
Annls Mus. r. Congo belge Sér. 8vo, 60: 95; 1958, Bull. Soc. r. Zool. Anvers,
9: 7; 1968, Bull. Annis Soc. r. ent. Belg., 99: 168. Neonyssus (Paraneonyssus)
emberizae, Bregetova, 1967, Parazit. Sb., 23: 133.
Previous records (both Passeriformes).—Common swallow (vagrant),
Hirundo rustica Linnaeus (Hirundinidae). Gouldian finch, Poephila gouldiae
(Gould) (Ploceidae).
Fain (1963d) comments on the close relationship between this species
and P. icteridius (Strandtmann and Furman, 1956). Should they prove
Synonymous, the former has priority by six months.
340° THH NASAL MITES OF QUEENSLAND BIRDS
| Figs 82-89. Ptilonyssus capitatus (Strandtmann) (@Q from Mirafra javanica) —82-83,
Lateral views of tarsus IV. 84-85, Dorsal and ventral views of tarsus II. 86, Dorsal
view of tarsus I. 87, Ventral view of capitulum (left palp in dorsal view). 88-89, Dorsal
and ventral views of idiosoma, with anal shield foreshortened.
ROBURT DOMROW 341
PTILONYSSUS PYGMAbUS (Bregetova), n. comb.
Neonyssus (Spizonyssus) pygmacus Bregetova, 1967, Parazit. Sb., 23: 131.
Previous record.—Goldfinch (introduced), Carduelis carduclis (Linnaeus)
(Fringillidae, Passeriformes) .
PTILYONYSSUS HIRSTI (de Castro and Pereira)
Neonyssus hirsti de Castro and Pereira, 1947, Archos Inst. biol., S Paulo,
18:129; Porter and Strandtmann, 1952, Tex. J. Sci., 4: 394. Paraneonyssus
hirsti, Feider, 1962, Studi Cere. stint. Iasi, 13:48. Ptilonyssus hirsti, Fain,
1963, Bull. Annls Soc. vr. ent. Belg., 99:170; Domrow, 1964, Acarologia,
6: 608. Ptilonyssus nudus Berlese, 1892, “Acari, myriapoda, et scorpiones
hucusque in Italia reperta, ordo Mesostigmata” (Patavii), fase. 54, No. 1 ( 2 ),
nec Berlese and Trouessart, 1889, Bull. Biblioth. scient. Ouest, 2: 130.
Ptilonyssus nudus Hirst, 1916, J. zool. Res., 1:78 (2), nec Berlese and
Trouessart, Loc. cit.
Previous record.—House-sparrow (introduced), Passer domesticus
(Linnaeus) (Fringillidae, Passeriformes), Brisbane.
This material compares well with specimens from the United States.
PTILONYSSUS NEOCHMIAR, Nn. Sp.
(Figs 78-80, 90-91)
Diagnosis According to Thomson (1964), Vidua Cuvier, Hypochera
Bonaparte, and Steganura Reichenbach are a group of three very similar
genus-group taxa of ploceid affinities, from which nasal mites of the genus
Ptilonyssus have been described as follows: P. viduae Fain (1956, 1957e),
P. hypocherae (Fain, 1963e), n. comb., and P. steganurae Fain (1967b).
P. viduae is the type-species of Passeronyssus Fain (1960f, 1962c), the
genus in which P. hypocherae was originally placed, and neither species shows
the stigmata extended anteriorly into peritremes. However, in P. steganurae,
and in the species with which Fain originally thought to associate it, P.
intermedius (Hirst, 19216) (see also Fain, 1960a; Fain and Hyland,
19626), peritremes are present.* Fain also compares P. steganurae with
P. ploceanus Fain (1956), a species he treats more fully in 1957e, together
with several other related species, all possessing peritremes, from other
ploceid and fringillid genera. It is clear, therefore, that we are in the presence
of congeners, and the essential criterion of difference listed by Fain between
Ptilonyssus and Passeronyssus (peritremes present or absent) no longer
holds good.
The new ploceid parasite, P. neochmiae, lacking peritremes, but possessing
an extensive opisthonotal shield, is therefore best compared, not with P.
viduae, but with P. hypocherae. It is readily separable, however, by the
absence of anterolateral extensions to the podonotal shield, the relatively
broader dimensions of the opisthonotal shield, and the length and position
of the anal setae.
*The unique specimen of P. intermedius is alleged on Trouessart’s label to have
come from a Malagasy bird, but it should always be kept in mind that, as this
worker collected much of his material from dried museum skins, his data should
always be verified by fresh material. For example, listrophorid mites said by Troues-
sart to have come from an African rodent, and described by Lawrence (1956) as
Cricetomysia andrei, proved to be Campylochirus chelopus Trouessart (1893), a parasite
of a peculiarly Tasmanian marsupial (Domrow, 1958). Also, Trouessart himself (1896)
expressed the probability that Chirodiscus amplexans Trouessart and Neumann (1890),
described from the feathers of an Australian bird, will also prove to belong to the
Listrophoridae, a family modified towards a lifelong attachment to single mammalian
hairs.
342 THE NASAL MITES OF QUEENSLAND BIRDS
Types.—Holotype female and four paratype females from a crimson finch,
Neochmia phaeton (Hombron and Jacquinot) (Ploceidae, Passeriformes) ,
Innisfail, 3.viii.1966, M. L. F. Holotype N. I. C.; paratypes R. D. and A. F.
Female.—Idiosoma 510 long in.slghtly distorted specimen figured.
Podonotal shield (Fig. 90) slightly longer than wide, 2380-240,» long, 210-—220yu
Figs 90-91. Ptilonyssus neochmiae, n. sp. (Q from Neochmia phaeton).—Dorsal and
ventral views of idiosoma.
Figs 92-96. Ptilonyssus sittae Fain (protonymph from Climacteris picumnus) —92-93,
Dorsal and ventral views of podosoma. 94, Anal shield. 95, Pygidium. 96, Ventral
view of capitulum (right palp in dorsal view).
wide; anterior margin irregularly convex, sides subparallel, and posterior
margin straight. Setae in eight pairs; surface reticulate, strongly punctate,
and marked by muscle insertions. Opisthonotal shield sub-triangular, with
rounded corners, length 192-196n, width 187-192. Six setae normally present
anteriorly, in hexagonal arrangement; pygidial setae present. Dorsal cuticle
with three pairs of setae flanking podonotal, and several setae and pores
flanking opisthonotal shield. Stigmata without peritremes, borne on weakly
sclerotized, posteriorly extended stigmatic shields.
ROBERT DOMROW 343
Sternal shield (Fig. 91) very weakly sclerotized, with SI-III, but no
pores, on shield proper. Metasternal setae absent. Genital shield short and
broad, reticulate and shagreened, and with narrow, rayed operculum. Genital
setae set well forward on shield, with associated pores set free in cuticle
near posterior margin of shield. Anal shield angularly convex anteriorly,
and with cribrum posteriorly. Anus centrally placed, preceded by adanal,
and followed by postanal seta. Ventral cuticle with several pairs of setae
and pores. All body setae minute.
All leg setae minute, except for those at apex of tarsi IJ-IV (Figs
79-80). Coxae 2.2.2.1. Trochanter I (1-0/2-1), II (1-0/3-1), III-IV (1-0/3-0).
Femora I-II (1-4/1-1), III (1-38/1-0), IV (1-2/1-0) ; genua I-III (1-4/2-1),
IV (1-4(3)/1-0). Tibiae (1-8/2-1). Tarsi —.17.17.17 (mv absent). Setae
a1, avi, pvi, and pl, on tarsi II-IV, particularly ventral pair, considerably
stronger than remainder of setae on ventral aspect. Claws I weak, but
not clearly visible; ambulacrum I also weaker and more slender than II-IV.
Basis capituli (Fig. 78) with pair of weak capitular setae, but
deutosternum absent. HI absent, II-III present, but very weak. Palpal
setae weak, apart from two dorsodistal tibial rods. Tarsus obsolescent, with
two slender setae. Strong claw present. Chelicerae attenuate in distal half,
with chelate portion occupying one-thirteenth of total length. Tritosternum
absent.
PTILONYSSUS STRUTHIDHAE, N. Sp.
(Figs 97-106)
Diagnosis.—Leach (1958), whose classification of Australian birds I
am presently using, places Struthidea Gould and Corcoraxz Lesson among the
Corvidae and Grallina Vieillot as the sole member of the Grallinidae. However,
Mayr and Amadon (1951) have grouped these three genera — mainly on a
characteristic habit, all build a bowl-shaped mud nest — into the Grallinidae,
recognizing two subfamilies, Grallinae (sic) for the peculiar Grallina and
Corcoracinae for the more closely related Struthidea and Corcoraz.
This subfamilial classification parallels that of their respective nasal
mites — the slender-bodied Ptilonyssus grallinae Domrow (1964¢) is quite
distinct from the thick-set P. struthideae, n. sp., and P. corcoracis, n. Ssfp.,
in showing the following characters: podonotal shield strongly excavated
posterolaterally, thereby flanking peritrematalia; pygidial shield distinct;
anal shield elongate, with distinct cribrum, but without postanal seta; body
setae strong and peg-like; coxa IJ with anterodorsal process; leg chaetotaxy:
coxae 2.2.2.1; trochanters I-II (0-0/3-1), III (2-0/2-0), IV (1-0/5-0) ;
femur I (2-4/0-2), II (1-4/1-1), TI (1-8/1-0), IV (1-8/2-0); genua
I-III (1-4/0-1), IV (1-8/0-0); tibiae (1-8/2-1); tarsi -—.17.17.17 (mw
lacking) ; palpal chaetotaxy 1.2.4.6.
However, both in general (facies) and in particular (chaetotaxy of
legs and palpi), the two new species are very similar indeed and are separable,
as noted above, only on the degree of sclerotization of the dorsal shields
and the setation of tarsus IV.
In view of the uncertain relationships of their two host genera, it is
difficult to point to any species of Ptilonyssus as a possible relative, even
confining one’s attention to the species with extensive opisthonotal shields.
Of the Australian species in this category — omitting P. hirsti (de Castro
and Pereira, 1947) from the intreduced Passer domesticus, a fringillid — P.
ailuroedi Domrow (19646), from ptilonorhynchids, is readily separable by
its four immense mesonotal setae; P. colluricinclae Domrow (1964c), from
pachycephalids, by the shape and setation of the dorsal shields and the
armature of tarsi II-IV; and P. maluri Domrow (1965c), from a sylviid,
by the presence of metasternal setae and a distinct deutosternum.
THE NASAL MITES OF QUEENSLAND BIRDS
)
(20
oa
olor Meee 225
Gr Se .
° Q ro
ag BS 8
92 O
> )
eS Qo ww
2
2 | 106
| eee F ce
ge eee a (Sa | RD
Figs 97-106. Ptilonyssus struthideae, 0. sp. (9 from Struthidea cinerea) —97-99, Dorsal
and ventral views of idiosoma. 100-101, Dorsal and ventral views of leg IV. 102-103,
105, Dorsal and ventral views of tarsus IV.
Ventral and dorsal views of tarsus I. 104—
106, Ventral view of capitulum (left palp in dorsal view).
ROBDRT DOMROW 345
It has since come to my notice that P. struthideae is much more closely
related to P. nucifragae (Hirst, 1923) (see also Fain, 19600; Bregetova,
1967), whose host, Nucifraga caryocatactes (Linnaeus) is likewise a corvid
(Thomson, 1964), a relationship which would urge caution in accepting
Mayr and Amadon’s findings (1951). I have seen two pairs of Hirst’s
species, but the new species is readily distinguished by the different shapes
of both its dorsal shields, the posterior reduction of its anal shield, and
the uniformity in length of its coxal setae.
T'ypes.—Holotype female and two paratype females from apostle-birds,
Struthidea cinerea Gould. (Corvidae, Passeriformes), Condamine, 29.v.1965,
H. A. S.; Condamine, 6.vii.19638, I. D. F.; and Charleville, 1.11.1967, R. D.
and J.S. W. Holotype N. 1. C.; paratypes R. D.
Female.—Idiosoma relatively stout, 660» long in somewhat flattened
specimen. Podonotal shield (Figs 97-98) very slightly wider than long,
strongly arched anterolaterally, and virtually straight posteriorly; bearing
six Or seven pairs of minute setae. Opisthonotal shield one-third again
as long as wide, tapering irregularly to convex posterior margin; bearing
two pairs of setae anteriorly, and three to five irregularly arranged setae
posteriorly, none of which appear to be pygidials. Both shields closely
punctate except for very narrow marginal strip, marked by distinct muscle
insertions, and showing few weak pores. Body cuticle with four shieldlets
and six setae flanking podonotal, and about eight setae and pores flanking
opisthonotal. shield. Stigmata provided with short peritremes.
Sternal shield (Fig. 99) narrower anteriorly, virtually textureless, and
outlined only by cessation of cuticular striae; bearing SI-III, but without
pores. Metasternal setae absent. Genital shield short and broad, showing
few muscle insertions and reticulate pattern more densely sclerotized than
remainder of shield; with rayed operculum anteriorly and two minute setae
posteriorly. Anal shield weakly sclerotized, with anus almost at posterior
margin; adanal setae near anterior of anus, postanal submarginal; cribrum
quite absent. Cuticle with about six pairs of setae and few pores.
Legs stout, with truncate tarsi bearing ambulacra ventrodistally (Figs
100-101). Setae generally short and spinose. Coxae 2.2.2.1. Trochanter
I (1-0/2-1), II (0—-0/3-1), IIT (1-0/3-0), IV (1-0/2-0). Femur I (1-4/2-1),
IT (1-4/1-1), I1I-IV (1-3/1-0). Genu I (1-4/2-1), I-III (1-4/0-1), IV
(1-3/0-0). ‘Tibiae (1-3/2-1). Tarsi -.17.17.16 (II-IV lacking mv; IV also
lacking ads, its relative position in II and III being indicated in Figs 100
and 104 by a small circle; av, and pv, slightly stronger). Ambulacra [I-IV
(Figs 104-105) with strong articulatory sclerites supporting heavy claws.
Ambulacrum I similar (Figs 102-103), but much smaller.
Tritosternum and deutosternum absent (Fig. 106). Capitular and
hypostomal setae III minute; remaining two pairs of hypostomals obsolescent
or absent. Palpi with usual five segments, but tarsus much reduced and
completely obscured dorsally by tibia. Setal formula 0.2.4.6. Tarsus with
one smaller and two larger setae; claw present, with indication of bifid tip
under oil immersion. Cheliceral shaft of uniform diameter basally, but tapering
in distal third; chelatae portion occupying one-fifteenth of total length.
PTILONYSSUS CORCORACIS, N. Sp.
(Figs 107-116)
Diagnosis.—See that for P. struthideae, n. sp., above.
Types.—Holotype female from a _ white-winged chough, Corcorar
melanorhamphus (Vieillot) (Corvidae, Passeriformes), Condamine, 6.vii.1963,
) I. D. F. Holotype N. I. C.
E
346 THE NASAL MITES OF QUEENSLAND BIRDS
Also 62 2 and 26 6 from C. melanorhamphus, Charleville, 28.1.1967,
R. D. and J. 8S. W.
Female.—Idiosoma about 880 long in mounted, somewhat compressed
material. Podonotal shield (Fig. 11Q) approximately as wide as long, but
Figs 107-116. Ptilonyssus corcoracis, n. sp. (2 from Corcoraz melanorhamphus) —107,
Sternal shield and coxa IV. 108, Anal shield. 109, Sternal setae. 110, Podonotum, with
one mesonotal shieldlet. 111, Opisthonotum, with other mesonotal shieldlet. 112, Ventral
view of capitulum, with basis not clear (right palp in dorsal view). 113-114. Ventral
and dorsal views of leg III. 115-116, Dorsal and ventral views or tarsus I.
ROBERT DOMROW 347
narrower anteriorly and slightly convex midposteriorly; outline generally
indistinct and eroded, particularly around anterior and posterolateral muscle
insertions; bearing six pairs of minute setae and two pores; surface
granulate, marked by regular pattern of muscle insertions. Mesonotum with
two shieldlets. Posteromedially an irregular, asetose shield (Fig. 111)
flanked on each side by seta and pore: this possibly represents the pygidial
complex, although further to each side—and, I fancy, behind—there are
additional shieldlets. Body cuticle with occasional setae and pores whose
pattern is indiscernable. Stigmata provided with short peritremes.
Sternal plate (Fig. 109) obsolescent, flanked by short, spinose SI-ITT.
Metasternal setae absent. Genital shield (Fig. 107) as in P. struthideae, n. sp.
Anal shield (Fig 108) subcireular, very weakly sclerotized, without cribrum;
anus centrally placed, flanked by three weak anal setae.
Leg chaetotaxy (Figs 113-114) as in P. struthideae, but genu IV
(1-4/0-0) on one side of holotype, and tarsi —.17.17.17 (mv absent; adz
present on ITV). Tarsus I is depicted in Figs 115-116.
Basis capituli and hypostome obscured in holotype, but other specimens
show deutosternum absent, weak capitular setae present, and hypostomal]
setae (particularly I) obsolescent or absent (Fig. 112). Dorsodistal rods
on palpal tibia weaker than in P. struthideae. Chelate portion occupying
one-twentieth of total length. Tritosternum absent.
PTILONYSSUS TRISCUTATUS (Vitzthum)
Ptilonyssoides triscutatus Vitzthum, 1935, J. Orn., Lpz., 83: 581.
Ptilonyssus triscutatus, Fain, 1957, Annls Mus. r. Congo belge Sér. 8vo, 60:
121; Domrow, 1966, Proc. Linn. Soc. N.S.W., 90: 196.
Previous record.—Rainbow-bird, Merops ornatus Latham (Meropidae,
Coraciiformes), Esk. Also Kowanyama.
PTILONYSSUS DIOPTRORNIS Fain
Ptilonyssus dioptrornis Fain, 1956, Revue Zool. Bot. afr., 53: 137; 1957,
Annls Mus. r. Congo belge Sér. 8vo, 60: 124; 1959, J. ent. Soc. sth. Afr.,
22: 26. Passeronyssus dioptrornis, Fain, 1960, Revue Zool. Bot. afr., 61: 110;
1962, Ibid., 66: 139; Nadchatram, McClure, and Lim, 1964, J. fed. Malay St.
Mus., 9: 105. Ptilonyssoides dioptrornis, Strandtmann, 1960, J. Kans. ent.
Soc., 833: 142. Passeronyssus enicuri Fain and Nadchatram, 1962, Bull. Annls
Soe. r. ent. Belg., 98: 276. New synonymy. Passeronyssus lusciniae Fain,
1962, Revue Zool. Bot. afr., 66: 140. New synonymy.
This species may now be recorded from Australia: 112 2, 14, 2
deutonymphs (one enclosing well developed ?), and 4 protonymphs from
pale-yellow robins, EHopsaltria capito Gould (Muscicapidae, Passeriformes),
Innisfail, 6.vii and 1l.viii.1965, G. J. B. and H. I. McD., and Wilson’s Peak,
15.v.1967, RD: and B: H. K.*
The three nominal species comprising this parasite of Turdidae and
Muscicapidae (two groups often considered merely as muscicapid subfamilies)
do not differ in any way that I would consider beyond the range of individual
variation. I have noted above on several occasions that many recent taxa,
both at species- and genus-group levels, have been erected with only minor
differences in mind. It is convenient here to note that it is perhaps even
more important to keep similarities in mind.
* The type-host of P. lusciniae, the nightingale, Luscinia megarhynchos Brehm
(Turdidae), was once introduced into Australia, but failed to become established
_ (Cayley, 1963).
348 THE NASAL MITES OF QUEENSLAND BIRDS
PTILONYSSUS DIcRURI Fain
Ptilonyssus dicruri Fain, 1956, Revue Zool. Bot. afr., 53: 137; 1957,
Annls Mus. r. Congo belge Sér. 8vo, 60: 122. Ptilonyssoides dicruri,
Strandtmann, 1960, J. Kans. ent. Soc., 33: 140. Passeronyssus dicruri,
Domrow, 1965, Acarologia, 7: 451. Ptilonyssoides faini Strandtmann, 1960,
J. Kans. ent. Soc., 33: 142.
Previous record.— Spangled drongo, Chibia bracteata (Gould)
(Dicruridae, Passeriformes), Samford and Esk. Also Kowanyama.
J have examined one Thai specimen identified as P. dicruri by Strandtmann.
PTILONYSSUS sITTAE Fain
(Figs 92-96, 219-222)
Ptilonyssus sittae Fain, 1965, Revue Zool. Bot. afr., 72: 158; 1966,
Bull. Annls Soc. r. ent. Belg., 102: 117.
This species, originally described from a European species of Sitta
Linnaeus (Sittidae), may now be recorded from Australia: 1 protonymph
from a brown tree-creeper, Climacteris picumnus Temminck (Certhiidae,
Passeriformes), Winbin Creek, 20.1.1966, R. D., D. J. M., and J. S. W.
Keast (1957) placed Climacteris Temminck in the family Sittidae, and
the matter is further discussed by Thomson (1964).
PTILONYSSUS AILUROEDI Domrow
Ptilonyssus ailuroedi Domrow, 1964, Acarologia, 6: 31, 619; Wilson,
1964, Pacif. Insects, 6: 381.
Previous record.—Green catbird, Ailuroedus crassirostris (Paykull)
(Ptilonorhynchidae, Passeriformes), Mt. Glorious. Also Wilson’s Peak.
Also 12 and 1 ? (in poor condition) from A. crassirostris (listed on the
label under its synonym A. smithi (Vigors and Horsfield)), Tweed River,
N.S.W., 11.1892, T. S. (see comments above on Tinaminyssus trichoglossi).
PTILONYSSUS ELBELI (Strandtmann), n. comb.
Cas elbeli Strandtmann, 1960, J. Kans. ent. Soc., 33: 1387. Sternostoma
bruxellarum Fain, 1961, Bull. Annls Soc. r. ent. Belg., 97: 53. New synonymy.
Previous records (both Sturnidae, Passeriformes).—European starling
(introduced), Sturnus vulgaris Linnaeus. Common myna _ (introduced),
Acridotheres tristis (Linnaeus).
I have examined one of Strandtmann’s paratypes.
PTILONYSSUS ANGRENSIS (de Castro), n. comb.
Rhinonyssus (Rhinacarus) angrensis de Castro, 1948, Archos Inst. biol.,
S Paulo, 18: 257. Cas angrensis, Baker and Wharton, 1952, “An introduction
to acarology” (Macmillan: New York), p. 81; Strandtmann and Wharton,
1958, Contr. Inst. Acar. Unw. Md, 4: 169.
This is the first record of this parasite of swallows outside the Americas:
722 from the welcome swallow, Hirundo neoxena Gould (Hirundinidae,
Passeriformes), Kowanyama, 21.x.1966, R. D. and H. A. S. They compare
well with 19from the United States.
PTILONYSSUS NITZSCHI (Giebel)
Dermanyssus Niteschi Giebel, 1871, Z. ges. Naturw., 38: 32. The original
capitalized spelling should be corrected under Art. 32. Rhinonyssus niteschi,
Vitzthum, 1935, J. Orn., Lpz., 88: 572 (pro parte). Neonyssus (Vitenyssus)
ROBDRT DOMROW 349
nitechi (sic) de Castro, 1948, Archos Inst. biol., S Paulo, 18: 277. Ptilonyssus
(Rhinonyssoides) nitschi (sic) Pereira and de Castro, 1949, Archos Inst.
biol., S Paulo, 19: 222. Vitenyssus niteschi, Strandtmann and Wharton,
1958, Contr. Inst. Acar. Univ. Md, 4: 175; Strandtmann, 1960, 7. Kans. ent.
Soc., 38: 131. Astridiella caprimulgi Fain, 1957, Riv. Parassit., 18: 96.
Previous record.—lLarge-tailed nightjar, Caprimulgus macrurus Horsfield
(Caprimulgidae, Caprimulgiformes) .
I agree with Strandtmann and Wharton (1958) that (7) it is unnecessary
to ‘delete D. niteschi, the first nasal mite described from birds, from the
nomenclature; and (i) it would be legitimate to consider the holotype of
A. caprimulgi as the neotype of D. niteschi. See discussion on Boydaia
crassipes (Berlese and Trouessart) below.
It is noteworthy that two of the hosts of P. mitzschi listed by Fain
(1957e) also harbour P. scotornis Fain (1956, 19576). However, the proportions
of the anal shield and the chelicerae appear to distinguish the two taxa. It
was, however, illegal of Fain to discard his original holotype of P. scotornis
for a better specimen from a different host.
PTILONYSSUS ECHINATUS Berlese and Trouessart
Ptilonyssus echinatus Berlese and Trouessart, 1889, Bull. Biblioth. scient.
Ouest, 2: 129; Strandtmann, 1960, J. Kans. ent. Soc., 33: 146; George, 1961,
J. Kans. ent. Soc., 34: 114; Domrow, 1964, Acarologia, 6: 607. Hapalognatha
platytricha Butenko, 1959, Nauwch. Dokl. vyssh. Shk., 2: 17 (nomen nudum) ;
1960, Zool. Zh., 39: 1494. Ptilonyssus chalybeaedomesticae do Amaral, 1967,
Archos Inst. biol., S Paulo, 34: 169. New synonymy. Ptilonyssus echinatus
taperaefuscae do Amaral. 1967. Archos Inst. biol., S Paulo, 34: 199. New
synonymy.
Previous records (both Hirundinidae, Passeriformes ).—Welcome swallow,
Hirundo neoxena Gould, Brisbane. Also Kowanyama. Common swallow
(vagrant), H. rustica Linnaeus.
New host record.—Tree-martin, Hylochelidon nigricans (Vieillot)
(Hirundinidae), Charleville, 31.1.1967, R. D. and J. S. W. (12).
All Australian specimens show the pygidial shield divided, as does 19
from H. rustica from the United States.
The above taxa would seem to provide another example of clinal
variation. All are parasites of swallows.
PTILONYSSUS PITTAE Domrow
Ptilonyssus pittae Domrow, 1964, Acarologia, 6: 26, 618.
Previous record.—Noisy pitta, Pitta versicolor Swainson (Pittidae,
Passeriformes), Upper Brookfield. Also Innisfail.
PrILONYSsuUS cRAcTIcI Domrow
(Figs 117-126)
Ptilonyssus cractici Domrow, 1964, Acarologia, 6: 598, 616; 1966, Proc.
Linn. Soc. N.S.W., 90: 196.
Previous records—Uaughing kookaburra, Dacelo gigas (Boddaert)
(Alcedinidae, Coraciiformes), Esk. White-winged triller, Lalage tricolor
(Swainson) (Campophagidae, Passeriformes), Kowanyama. Also Charleville.
Black-backed magpie, Gymnorhina tibicen (Latham) (Cracticidae, Passeri-
formes), Brisbane and Condamine. Also Esk. Pied butcher-bird, Cracticus
350 THE NASAL MITES OF QUEENSLAND BIRDS
nigrogularis (Gould), Logan Village. Grey butcher-bird, C. torquatus
(Latham), Esk. Black-backed butcher-bird, C. mentalis Salvadori and
d’Albertis, Chillagoe.
New host record.—Pied currawong, Streptera graculina (Shaw)
(Cracticidae), Esk, 10.vii.1967, R. D. and B. H. K. (7 2 2).
The leg chaetotaxy of specimens from cracticids is as follows. Coxae
2.2.2.1. Trochanters 4.5.4.4. Femur I (2-4/1-2), IT (1-4/1-1), HLIV
(1-8/0-0). Genua I-III (1-4/0-1), IV (1-2/0-0). Tibiae I-II (1-3/2-1),
III-IV (1-3/2-0). Tarsi -.17.17.17.. The two posterodorsal setae on genu
118 ye
yas
C20
Figs 117-118. Ptilonyssus cractici Domrow. (¢ from Lalage tricolor).—Dorsal and ventral
views of idiosoma.
III are set in relatively larger alveoli. Specimens from Lalage Robineau—
Desvoidy differ only in showing trochanter II 4. The specimen from Dacelo
shows a different chaetotaxy on two segments, viz. femur IV (1-8/1—0) and
tibia IV (1-8/2-1). See also note below on P. sphecotheris Domrow.
One magpie in subadult plumage (Esk, 13.11.1968, R. D. and B. H. K.)
yielded 105 9 2,9 8 8, 7 deuto-, and 22 protonymphs. This extremely heavy
infestation was confined to the nasal passages, the mites in the drier anterior
portions near the nostrils being partially encased in dried nasal secretions.
No mites were found in the trachea or lungs.
ROBHRT DOMROW 351
a
i
iA
‘ih
a
iN
if
Figs 119-121. Ptilonyssus cractici Domrow (Q from Lalage tricolor) —119-120, Dorsal
and ventral views of leg IV. 121, Ventral view of capitulum (left palp in dorsal view).
Figs 122-126. Ptilonyssus cractici Domrow (2 from Dacelo gigas) .—122-123, Dorsal and
ventral views of idiosoma. 124, Ventral view of capitulum (with left palp in dorsal
view). 125-126, Dorsal and ventral views of leg IV.
352 THE NASAL MITES OF QUEENSLAND BIRDS
PTILONYSSUS MOTACILLAE Fain
(Figs 127-132)
Ptilonyssus motacillae Fain, 1956, Revue Zool. Bot. afr., 53: 143; 1957,
Annls Mus r. Congo belge Sér. Svo, 60: 104, 187; 1959, J. ent. Soc. sth. Afr.,
22: 24; 1960, Revue Zool. Bot. afr., 61: 109; 1962, Lbid.. 66: 128; Domrow,
1964, Acarologia, 6: 600; 1965, Ibid., 7:460. Ptilonyssus motacilla (sic)
Gretillat, 1961, Vie Milieu, 12: 155. Ptilonyssus estrildicola Fain, 1959, J.
ent. Soc. sth. Afr., 22:22. New synonymy. Ptilonyssus fringilliicola Fain,
1959, J. ent. Soc. sth. Afr., 22: 28;,1962, Bull. Annls Soc. r. ent. Belg., 98:
262. New synonymy. Hapalognatha prima Butenko, 1959, Nauch. Dokl.
vyssh. Shk., 2:17 (nomen nudum) ; 1960, Zool. Zh., 39: 1490. The two species
headings on this latter page are transposed. Ptilonyssus lobatus Strandtmann,
1960, J. Kans. ent. Soc., 33: 147. New synonymy. Ptilonyssus cinnyricinch
Fain, 1962, Revue Zool. Bot. afr., 66: 132. New synonymy. Ptilonyssus
estrildicola taeniopygiae Fain, 1963, Revue Zool. Bot. afr., 68: 74; Nadchatram,
McClure, and Lim, 1964, J. fed. Malay St. Mus., 9: 105. New synonymy.
Ptilonyssus motacillae phoenicuri Fain, 1966, Revue Zool. Bot. afr., 74: 90.
New synonymy. Ptilonyssus sp. Fain and Nadchatram, 1962, Bull. Annls
Soc. r. ent. Belg., 98: 280. See also Domrow (19660).
Previous records (all Passeriformes).—Rufous whistler, Pachycephala
rufiwentris (Latham) (Pachycephalidae), Mt. Cotton and Samford. Also
Esk and Kowanyama. Australian pipit, Anthus australis Vieillot (Mota-
cillidae), Esk. Yellow wagtail (vagrant), Motacilla flava Linnaeus. Goldfinch
(introduced), Carduelis carduelis (Linnaeus) (Fringillidae). Zebra finch,
Taeniopygia castanotis (Gould) (Ploceidae). Common myna (introduced),
Acridotheres tristis (Linnaeus) (Sturnidae).
New host records (both Passeriformes).—Rose-robin, Petroica rosea
Gould (Muscicapidae, Passeriformes), Esk, 17.iv.1967, R. D. and B. H. K.
(4 92, 1 deutonymph). Spice finch (introduced), Lonchura punctulata
(Linnaeus) (Ploceidae), Innisfail, 3.viii.1965, R. D. and J. S. W. (12).
Fain (1962c) stated that “P. motacillae présente des légéres variations
en rapport avec la nature de Vhéte parasité. Ces variations portent sur la
forme et les dimensions des écussons et des poils dorsaus, les dimensions du
corps et des différents organes, la structure de Vécusson pygidial (simple ou
double). Comme il existe tous les intermédiaires entre les formes extrémes
il ne semble pas indiqué de donner a ces variations un rang spécifique ni méme
subspécifique”’. Later (19666), however, he was to describe P. m. phoenicuri.
A further Australian variant from Petroica Swainson is figured on page 353.
In the former paper, Fain also described P. cinnyricincli as intermediate
between P. motacillae and P. lobatus in showing incipient mesonotal lobes.
Weaker lobes are also present in P. m. phoenicuri and in Australian specimens
from Pachycephala Vigors. |
Fain (1959, 1962a) noted variation in the pygidial shield of specimens
of P. fringilliicola from Emberiza in Africa and Carduelis Brisson in Europe,
and one of the latter specimens is very similar to P. motacillae. In particular,
it shows a process anterodorsally on coxa II as well as ventrointernally on
the palpal trochanter. The African P. estrildicola and its Australian
subspecies, P. e. taeniopygiae, would also be synonymous by the present
criteria. The two-toned illustrations of P. tillae Fain (1959), from an
African starling, are too dense to allow of a decision on its status.
There has also been described (de Castro, 1948; George, 1961; Fain,
19646) a group of closely related taxa from neotropical and _ nearctic
passerines, but I have no experience of them. Their chefs de file are P. sairae
de Castro and P. japuibensis de Castro.
ROBERT DOMROW Bde
Figs 127-132. Ptilonyssus motacillae Fain (@ from Petroica rosea)— 127-128, Dorsal
and ventral views of idiosoma. 129, Ventral view of capitulum, with palpal tarsus
not clear (left palp in dorsal view). 130, Ventrolateral view of chelicera. 131-132,
Dorsal and ventral views of leg IV.
354 THH NASAL MITES OF QUEENSLAND BIRDS
The only tangible way in which the specimens from Petroica Swainson
differ from the earlier Australian material is that the pygidial shield is
entire rather than divided (Fig. 127).
Specimens from all three Australian hosts have the following leg
chaetotaxy. Coxae 2.2.2.1. Trochanters 4.4.4.5. Femur I (2-4/1-2),
(1-4/1-1), III (1-8/0-0), IV (1-8/1-0). Genua I-III (1-4/0-1), IV
(1-2/0-0). Tibiae I-IT (1-8/2-1), I1I-IV (1-3/2-0) but often (1-3/1-0).
Tarsi —.17.17.17 (mv absent).
Fain’s P. e. estrildicola came from various African ploceids, and his later
P. e. taeniopygiae from T. castanotis which died in Antwerp shortly after
their importation from Australia. In view of the large number of species of
various families of mites parasitizing the nasal passages of birds now known
to occur in related African, S.E. Asian, and Australian hosts, I would
accept this record as a natural infestation. The specimen from L. punctulata
shows the three anal setae in almost the same transverse line, cf. P. e.
taeniopygiae.
I have been able to examine 1? from Carduelis (labelled P. fringillicola)
and 6 2 @ paratypes of P. lobatus.
PTILONYSSUS LANGEI (Butenko), n. comb.
Hapalognatha langet Butenko, 1959, Nauch. Dokl. vyssh. Shk., 2: 17
(nomen nudum) ; 1960, Zool. Zh., 39: 1490. The two species headings on this
latter page are transposed.
Previous record.—Common swallow (vagrant), Hirundo rustica SULEEEOS
(Hirundinidae, Passeriformes).
PTILONYSSUS ORTHONYCHUS, N. Sp.
(Figs 133-137)
Diagnosis.—This morphologically nondescript species seems best compared
with P. malaysiae Fain (1964a@), originally described from the fairy bluebird,
Irena puella (Latham). Irena Horsfield, incidentally, is placed at the end
of the family Aegithinidae by McClure (1963), leading immediately into
the Timaliidae, to which Orthonya Temminck, the host of the new species,
belongs. However, in all the following characters, P. malaysiae is at variance
with P. orthonychus: podonotal shield associated with seven pairs of setae
(excluding five pairs set between the shield and the peritrematalia — these
are commonly present throughout the species of Ptilonyssus) ; genital setae
not set on shield; adanal setae set in front of anus; coxa IT with anterodorsal
process; tarsi II-IV with av; and pv; spur-like; capitular setae absent.
If the Timaliidae are considered merely a subfamily of the Muscicapidae |
(Keast, 1961; Thomson, 1964), the only Australian species recalling P.
orthona ychus ns) 12 microecae Domrow (19666). The two may, however, be
separated by setational details of the podonotal and anal shields, legs, and
capitulum.
Types.—Holotype female and eight paratype females from southern
chowchillas, Orthonyx temminckii Ranzani (Timaliidae, Passeriformes),
Wilson’s Peak, 15.v.1967, R. D. and B. H. K. Holotype N. I. C.; paratypes
R. D. and A. F.
Female.—A lightly sclerotized species with particularly weak setation
(Fig. 133); idiosoma 825p long in engorged, somewhat flattened specimen
figured. Podonotal shield suboval, but slightly broader and weakly concave
anteriorly, length 245-286, breadth 183—250u; associated setae in nine pairs,
of which verticals and one or two anterolateral pairs are normally free in
ROBERT DOMROW 355
adjacent cuticle. Cuticle between shield and peritremes with additional five
pairs of setae arranged 1.1.8. Mesonotum with usual four shieldlets and
transverse band of ten setae. Pygidial shield entire, transverse, with two
setae and accompanying pores, and surrounded by about five pairs of setae
slightly stronger than remainder of dorsal series.
Figs 133-137. Ptilonyssus orthonychus, n. sp. (2 from Orthonyx temminckii) —133-134,
Dorsal and ventral views of idiosoma. 135, Ventral view of capitulum (right palp
in dorsal view). 136-137, Dorsal and ventral views of leg IV.
Fig. 138. Sternostoma neosititae, n. sp. (2 from Neositta striata).—Dorsal view of
tarsus I.
Sternal shield (Fig. 134) evanescent, marked only by cessation of cuticular
striae, bearing four pores and SII, leaving SI and III free in cuticle. Meta-
sternal setae absent. Genital shield ligulate, with few indications of muscle
insertions and rayed operculum; genital setae set on shield, but attendant
pores free in cuticle. Anal shield elongate, strongly arched anteriorly; anus
356 THE NASAL MITES OF QUEENSLAND BIRDS
in front of all three anal setae; cribrum present. Ventral cuticle with about
ten pairs of slightly stronger setae.
Coxa II without anterodorgal process and leg setation undistinguished
(Figs 186 and 137). Coxae 2.2.2.1. :Trochanters I-II (1-0/2-1), ILI-IV
(1-0/3-0). Femur I (2-4/1-2), II (i1-4/1-1), III (1-8/0-0), IV (1-8/1-0).
Genua I-III (1-4/0-1), IV (1-3/0-0). Tibiae (1-8/2-1). Tarsi —.17.17.17
(mv absent; II-IV with av; and pv; only very slightly enlarged).
Basis capituli (Fig. 185) with two capitular setae and about seven
small denticles in single file in deutosternal groove. Hypostome with HIIT
subequal to capitular setae, but HI and II obsolescent. Palpal setal formula
1.2.4.8. Tarsus with several weak setae and remnant of claw. Chelicerae
attenuate in distal three-fifths, chelate portion occupying one-twentieth of
total length. Tritosternum absent.
PTILONYSSUS MICROECAE Domrow
Ptilonyssus microecae Domrow, 1966, Proc. Linn. Soc. N.S.W., 90:
196 ; 1967, Tbid., 91: 219.
Previous records.— Jacky winter, Microeca fascinans (Latham)
(Muscicapidae, Passeriformes), Esk. Also Charleville and Kowanyama. Lemon-
breasted flycatcher, M. flavigaster Gould, Kowanyama.
PTILONYSSUS TERPSIPHONEI Fain
(Figs 189-152)
Ptilonyssus terpsiphonei Fain, 1956, Revue Zool. Bot. afr., 53: 145; 1957,
Annls Mus. r. Congo belge Sér. 8vo, 60: 104, 187; 1959, J. ent. Soc. sth Afr.,
22: 22; Domrow, 1965, Acarologia, 7: 453.
Previous records (all Muscicapidae, Passeriformes).—Black-faced fly-
catcher, Monarcha melanopsis (Vieillot), Esk. Spectacled flycatcher, WU.
trivirgata (Temminck), Mt. Jukes. Also Innisfail. White-eared flycatcher,
Carterornis leucotis (Gould), Chelona. Also Innisfail.
New host records (all Muscicapidae).—Leaden flycatcher, Myiagra
rubecula (Latham), Esk, 6.x.1966, R. D. and J. S. W. (6 292). Restless
flycatcher, Seisura inquieta (Latham), Esk, 5.11966 and 17.iv.1967, R. D.,
B. H. K., and J. S. W. (42 2, 344, 1 protonymph). Australian- pied
flycatcher, Arses kaupi Gould, Innisfail,'5.viii.1965, R. D. and J. S. W. (12).
Shining flycatcher, Piezorhynchus alecto (Temminck), Innisfail, 3 and
6.vii1.1965, R. D. and J. 8. W. (8 2 29,4 ¢ ¢).
Individual variation in this material is considerable, but does not dispel
the conviction that only one species is involved. Fain (1957e) illustrated
three pairs of setae along the midposterior line of the podonotal shield in
his African material, and this is also the case in specimens from M. trivirgata
(Fig. 147). However, in all other Australian material, the anteriormost
pair is absent (except in one specimen from Seisura, where one seta of the
pair is present). Moreover, in specimens from Myiagra Vigors and Horsfield,
any one or two of the remaining four setae may be rudimentary or absent
(Fig. 139). Variation in the pygidial shield is indicated in Figs. 148-151.
Distinct poststigmatic shields are always present.
The sternal setae are normally filiform (Fig. 152), but SI are rod-like
in material from Seisura, and all are so modified in that from Myiagra (Fig.
140). The postanal seta is absent in specimens from M. melanopsis and
Carterornis Mathews, minute in those from Arses Lesson and Piezorhynchus
Gould, and otherwise normal.
ROBDRT DOMROW 357
144
wey dh VA
Figs 139-152. Ptilonyssus terpsiphonei Fain._139-140, Dorsal and ventral views of
idiosoma of 2 from Myiagra rubecula, with inset of pygidial shield from another
specimen from same host. 141, Dorsal and ventral views of genu-tibia III of 2 from
M. rubecula. 142-143, Dorsal and ventral views of genua-tibiae III-IV of 9° from
Monarcha trivirgata. 144-145, Dorsal and ventral views of leg IV of 2 from M. rubecula.
146, Ventral view of capitulum (right palp in dorsal view), with inset of chelicera,
both from @ from M. rubecula. 147, Podonotal shield of 2 from W. trivirgaia. 148-151,
Pygidial shield of 92 from Seisura inquieta, Arses kaupi, Piezorhynchus alecto, and
M. trivirgata. 152, Sternal shield of 2 from Carterornis leucotis.
358 THE NASAL MITES OF QUEENSLAND BIRDS
The legs present a mélange of rod-like and filiform setae (Figs 144-145).
Both setae on coxa I are always rod-like, and so is usually the anterior
seta on coxa II in material from Myiagra, Seisura, Piezorhynchus, and M.
trivirgata (Fig. 140). Otherwise, the coxal setae are filiform. The major
variant in the leg setation occurs on the genua of specimens from both
species of Monarcha Vigors and Horsfield: I (1-4/2-1), II-III (1-4/0-1),
IV (1-8/1-0), compared with I (1-2/2-1), II-III (1-2/0-1), IV (1-1/1-0)
in other specimens (Figs 141-143). I include the one specimen from Arses
in the latter category, although it shows I (1-38/2-1), IT (1-2/0-1), III
(1-2/1 av-1) and (1-2/1 pv-1), IV (1-1/1-0) and (1-1/2-0) — indeed, the
setation in this specimen varies on the four segments trochanter—tibia. The
femoral formulae vary slightly, but this is to be expected on the most setose
of the segments. An all-encompassing formula would be 10/9.8/7.5/6.5/6. The
remaining segments are constant: coxae 2.2.2.1; trochanters 4.4.4.5; tibiae
J-II (1-8/2-1), III-IV (1-8/2-0); tarsi —18.18.18 (mv present, see Fig.
145).
The capitular setae are normally present (Fig. 146), but appear to be
absent in material from M. melanopsis. The palpal formula (trochanter-genu)
is generally 0.2.2 (Fig. 146), but increases to 0.2/3.4 in specimens from
Monarcha and Carterornis.
PTILONYSSUS BRADYPTERI (Fain), n. comb.
Passeronyssus bradypteri Fain, 1962, Revue Zool. Bot. afr., 66: 148;
Domrow, 1966, Proc. Linn. Soc. N.S.W., 90: 196.
Previous record.—Rufous songlark, Cinclorhamphus mathewsi Iredale
(Sylviidae, Passeriformes), Esk.
PTILONYSSUS ACROCEPHALI Fain
(Figs 153-160)
Ptilonyssus acrocephali Fain, 1964, Bull. Annis Soc. r. ent. Belg., 100: 55.
This species, originally described from a European species of Acrocephalus
Neumann, may now be recorded from Australia: 5 9 @ and 1 protonymph
from Australian reed-warblers, A. australis (Gould )(Sylviidae, Passeri-
formes), Esk, 6.x.1966, and Kowanyama, 12.i11.1967, R. D., B. H. K., H. A. S.,
and J. S. W. The leg chaetotaxy (Figs 158-159) of these specimens is as
follows. Coxae 2.2.2.1. Trochanters 4.4.4.5. Femur I (2-4/1-2), II (1-4/1-1),
IlI-IV (1-4/1-0). Genua I-III (1-4/0-1), IV (1-8/0-0). Tibiae I-II
(1-3/2-1), ITI-IV (1-3/2-0). Tarsi —17.17.17 (mv absent). One more, or
one less seta was occasionally noted on trochanter IV, femora I-III, and
tibia III. The arrangement of setae on the capitulum is shown in Fig. 160.
The genital shield of Australian specimens (Fig. 154) takes in the genital
setae as in P. calamocichlae Fain (1956, 1957e), while these setae are shown
as free in Fain’s figure of P. acrocephali. Moreover, the degree of postero-
lateral erosion of the podonotal shield (Fig. 153) is such that the subposterior
pair of setae is discal, again as in P. calamocichlae, rather than marginal
as in P. acrocephali. The anterolateral margins of the shield are irregular
in Australian material (Figs 155-157). Actually, these two taxa are otherwise
very similar, and it may be that they are conspecific. At the moment,
however, the situation is clouded by several additional taxa recently described
from sylviids.
P. calamocichlae Fain (1956, 1957e) was originally described from five
African genera of Sylviidae: Calamocichla Sharpe, Hippolais Balderstein,
Cisticola Kaup, Prinia Horsfield, and Apalis Swainson. The material from
Hippolais has since been given subspecific status as P. c. hippolaisi by Fain
ROBHRT DOMROW 359
Ses 55
fi
RD
BHK
Figs 1538-160. Ptilonyssus acrocephali Fain (Q from Acrocephalus australis) —153-154,
Dorsal and ventral views of idiosoma. 155-157, Variations in setation of anterior
half of podonotal shield. 158-159, Ventral and dorsal views of leg IV. 160, Ventral
view of capitulum (left palp in dorsal view).
Figs 161-164. Ruandanyssus artami, n. sp (2 from Artamus cinereus) —161—162, Dorsal
and ventral views of tarsus I. 163-164, Dorsal and ventral views of leg IV.
360, THE NASAL MITES OF QUEENSLAND BIRDS
(1963e), while the material from the remaining three genera had earlier
been given specific rank as P. cisticolarum by Fain (1959). Finally, the
material from Prinia and Apalis, showing the pygidial shield divided rather
than entire as in the three preceding taxa, was given specific rank as
P. elongatus by Fain (1964d).
In the meantime, two further taxa with even more strongly divided
pygidial shields were described from Belgian sylviids by Fain (1962a, 1963d) :
P. phylloscopi from Phylloscopus Boie and P. rwandae sylviae from Sylvia
Scopoli.
Material from another Australian sylviid genus (Gerygone Gould) is
considered below.
PTILONYSSUS MONARCHAB, D. Sp.
(Figs 165-170)
Diagnosis.—Of the described species of Ptilonyssus from Australian
flycatchers, only two species, P. terpsiphonei Fain (1956, 1957e) and P.
microecae Domrow (19666), show the podonotal shield complete enough
posteriorly to take in the median pair of setae set immediataely in front of
the mesonotal shieldlets. This is also true of P. monarchae, which may best
be compared with P. microecae. However, while the anterior process on
coxa II is absent, and the postanal seta present in the former, the opposite
obtains in the latter. There are also minor differences in the setation of
the podonotal shield and the position of the adanal setae relative to the
anus.
Types.—Holotype female from a spectacled flycatcher, Monarcha trivirgata
(Temminck) (Muscicapidae, Passeriformes), Innisfail, 31.viii.1965, H. I. MeD.
Holotype N. I. C.
Female.—Idiosoma 670 long in unengorged condition, broader in anterior
half. Podonotal shield (Fig. 165) half again as long as wide, 256 x178p,
narrower in posterior third; anterior margin subrectilinear, posterior margin
convex; with muscle insertions and seven pairs of setae as indicated.
Peritremes short, borne on oval shieldlets with poststigmatic portions more
heavily sclerotized. Body cuticle between and around podonotal shield and
peritremes with 13 setae. Mesonotum with two shieldlets and transverse
band of ten setae. Pygidium with entire shield and eight pairs of setae.
Sternal shield (Fig. 166) evanescent, with six setae, but apparently
without pores. Metasternal setae absent. Genital shield drop-shaped, with
rayed operculum and two genital setae, the latter accompanied by pores
in adjacent cuticle. Anal shield elongate, with anus in front of adanal
setae; postanal seta absent, but cribrum present. Body cuticle with fifteen
setae arranged 2.5.8.
Coxa II with slender process on anterodorsal margin. Coxae 2.2.2.1.
Trochanters I-III (1-0/2-1), III (2-0/2-0), TV (1-1/3-0). Femur I (2-4/1-2),
IT (1-4/1-1), III (1-3/1-0), IV (1-8/2-0). Genua I-III (1-4/0-1), IV
(1-2/0-0). Tibiae (1-8/2-1). Tarsi -.17.17.17 (mv absent; I with one
ventrodistal seta stronger, II-IV with av; and pv; stronger, see Figs 167-168).
Basis capituli with two, and hypostome with six setae; HIT smaller,
remainder subequal (Fig. 170). Deutosternum with single file of ten denticles.
Palpal setal formula 1.3.4.8. Tarsus with about seven minute setae and bifid
claw. Chelicerae (Fig. 169) attenuate for slightly more than half their
length; digits occupying one-fifteenth of total length. Minute tritosternal
remnant present.
ROBERT DOMROW 361
Figs 165-170. Ptilonyssus gliciphilae Domrow (Q from Myzomela pectoralis) —165-166,
Dorsal and ventral views of idiosoma. 167-168, Dorsal and ventral views of leg IV.
169, Chelicera. 170, Ventral view of capitulum with inset showing tritosternum (left
palp in dorsal view).
Figs 171-173. Ptilonyssus gliciphilae Domrow (2 from Myzomela pectoralis) —171-172,
Dorsal and ventral views of opisthosoma. 173, Podonotal shield.
F
362: THH NASAL MITES OF QUEENSLAND BIRDS
PTILONYSSUS SETOSAH, 0. Sp.
(Figs 174-186)
Diagnosis.—The species of Ptilonyssus known from Rhipidura Vigors
and Horsfield (see Fain, 1968e, and Domrow, 19660) may be separated by
the following couplets:
il Vertical setae present, t.e. podonotal shield with seven pairs of setae.
Postanal’ ‘seta, presenti. Ae atic’ sere ote eins aes ead A ee ee
Vertical setae absent, i.e. podonotal shield Srila six pairs of setae (including
submarginal anterolateral pair). Postanal seta absent .... macclurei Fain
2 (1). Pygidial shield entire. Adanal setae in front of anus ........ setosae, nN. Sp.
Pygidial shield divided. Adanal setae behind anus .... rhipidurae Domrow
Types.—Holotype female and seven paratype females from a northern
fantail, Rhipidura setosa (Quoy and Gaimard) (Muscicapidae, Passeriformes),
Ella Bay, 4.viii.1965, R. D. and J. S. W. Holotype N. I. C.; paratypes R. D.
The following specimens are assigned to the new species, but do not
form part of the type series: 62 @ and 1 protonymph from rufous fantails,
R. rufifrons (Latham), Innisfail, 24 and 29.vi.1965, G. J. B. and H. I. MeD.
Female.—Idiosoma 594-616 long, 3224 wide in unfed specimens, and
up to 814-880 long, 385-418. wide in engorged material (all mounted).
Q
Figs 174-180. Ptilonyssus setosae, n. sp. (? from Rhipidura setosa).—174-175, Dorsal
and ventral views of idiosoma. 176-177, Variations in setation of podonotal shield.
178, Ventral view of capitulum (right palp in dorsal view). 179-180, Dorsal and
ventral views of leg IV.
ROBHRT DOMROW 363
Podonotal shield rather variable in shape, but generally slightly concave
anteriorly and posteriorly, with midlateral convexity; usually in range
178-201 long, 156-171 wide, but one specimen smaller, 169 *147p. Five
specimens show six pairs of setae on shield (Fig. 176), but I believe normal
complement to be seven pairs, attained only when midposterior arch of four
is full (Fig. 174). One specimen (Fig. 177) is grossly abnormal, both lacking
one anterolateral on each side and one median, as well as having midposterior
four irregularly arranged. Surface of shield weakly granulate and marked
by muscle insertions. Cuticle surrounding shield with six pairs of setae.
Peritremes present, with weak poststigmatic shields. Mesonotum with four
shieldlets and ten setae. Pygidial shield entire, subquadrate, with two
setae and some pores; surrounded by about eight setae. Dovwal setae in
form of stout spines.
Sternal shield delineated only by cessation of cuticular striae, without
evident texture (Fig. 175), bearing six setae and four pores submarginally.
Metasternal setae absent. Genital shield elongate, with longitudinal granula-
tions and weakly marked muscle insertions; operculum rayed. Genital
setae and accompanying pores free in cuticle. Anal shield broadest towards
front, with adanal setae in front of anus; postanal seta and cribrum present.
Ventral cuticle with about 10-12 pairs of spinose setae.
Legs provided with stout, spine-like setae ventrally, and much weaker
setae dorsally (Figs 179-180). Coxae 2.2.2.1 (on one side of one specimen
the seta on .coxa IV is bifid). Trochanters 4.4(5).4.5. Femur I (2-4/1-2),
II (1-4/1-1), Ill (1-8/1-0), IV (1-3/2-0). Genua I-III (1-4/0-1) but
I often (1-3/0-1), IV (1-2/0-0). Tibiae I-IT (1-8/2-1), ITI-IV (1-8/2-0).
Tarsi —17.17.17 (mv lacking). Ambulacrum I more elongate, and with
weaker, straighter claws than IT-IV.
Basis capituli and hypostome with four pairs of subequal, spinose setae
(Fig. 178). Deutosternum with about nine denticles in single file. Palpal
setal formula 1.2.4.7. Chelicerae suddenly attenuate in distal half, with
digits occupying one-twentieth of total length. Tritosternum absent.
Remarks.—The above description is based on specimens from R. setosa.
Those from R. rufifrons differ in the following respects. Idiosoma 737 x 330u
in moderately fed, 814-847, long and 352-3863» wide in engorged, and 880
374u in gravid specimens. Podonotal shield as long as wide, length 167-178,
breadth 174-1804; broadly arched anterolaterally and slightly concave
posteriorly. All dorsal setae in form of blunt rods (Fig. 181).
Venter with sternal and opisthosomal setae stouter (Fig. 182).
Setae on ventral aspect of legs in form of blunt rods (Figs 185-186).
Genua I and II often (1-8/0-1). Femur IV (1-3/1-0). In fact, all three
species included in the above key show the same gross setal formulae as
given by Domrow (19660) for P. rhipidurae, except that specimens of P.
setosae from R. setosa show an additional ventral seta on femur TV.
Capitulum also with rod-like setae on basis and hypostome (Fig. 183).
Chelicerae as in Fig. 184.
PTILONYSSUS GERYGONAE, Nl. Sp.
(Figs 190-194)
Ptilonyssus sp. Domrow, 1967, Proc. Linn. Soc. N.S.W., 91: 217.
Diagnosis—Of the taxa from sylviid genera diseussed above under
P. acrocephali Fain, P. gerygonae is most closely related to P. calamocichlae
hippolaisi Fain (1963e). However, the setae on the ventral surface of the
opisthosoma and in the transverse row of six immediataely behind the
. podonotal shield are undistinguished in the new species, while they are
364 THE NASAL MITES OF QUEENSLAND BIRDS
considerably enlarged in P. c. hippolaisi. Also, on the posterior tarsi, the
setae on the leading edge are retrorse, and av, and pv; divergent in P. ¢.
hippolaisi, but normally curved and with opposed tips, respectively, in the
new species. Finally, the bulbous basal portion of the chelicerae is relatively
longer in P. c. hippolaisit with respect to the attenuate distal portion.
Figs 181-186. Ptilonyssus setosae, n. sp. (? from Rhipidura rufifrons) —181-182, Dorsal
and ventral views of idiosoma. 183, Ventral view of capitulum (left palp in dorsal
view). 184, Chelicera. 185-186, Dorsal and ventral views of leg IV.
Figs 187-189. Ptilonyssus philemoni Domrow (@ from WMeliphaga notata) —187-188,
Ventral and dorsal views of opisthosoma. 189, Podonotal shield.
ROBPRT DOMROW 365
T'ypes.—Holotype female, six paratype females, and one protonymph from
black-throated warblers, Gerygone palpebrosa Wallace (Sylviidae, Passeri-
formes), Innisfail, 1.vii. and 16.viii.1965, G. J. B., R. D., H. I. McD., and
J.S. W. Holotype N. I. C.; paratypes R. D.
Female.—Idiosoma 704-836 long, 253-319. wide depending on degree
of engorgement. Podonotal shield (Fig. 190) 150-161» long, 138-147» wide;
broadly arched anterolaterally, irregularly bilobed posteriorly; bearing six
Figs 190-194. Ptilonyssus gerygonae, n. sp. (2 from Gerygone palpebrosa) —190-191,
Dorsal and ventral views of idiosoma. 192, Ventral view of capitulum (right palp in
dorsal view) (note tritosternum). 193-194, Dorsal and ventral views of leg IV.
pairs of setae (including vertical pair); surface weakly granulate and
marked by muscle insertions. One pair of setae immediately behind shield,
and five pairs arranged 1.1.3 between shield and peritrematalia. Mesonotum
with usual four shieldlets and transverse band of ten setae. Pygidial shield
entire, transverse, with two setae and pores; surrounded by about eight
setae. All dorsal setae in form of slender spines.
Sternal shield weakly outlined and textureless, with two pores; SI-III
in adjacent cuticle (Fig. 191). Metasternal setae absent. Genital shield
narrowly drop-shaped, leaving genital setae and attendant pores free in
366 THE NASAL MITES OF QUEENSLAND BIRDS
cuticle. Anal shield small, elongate, with adanal setae in front of, and postanal
seta behind anus; cribrum present. Ventral cuticle with about six pairs
of setae.
Legs with slender, spine-like sefae ventrally (Figs 193-194). Coxae
2.2.2.1. Trochanters 4.4.4.5. Femur I (2-4/1-2), II (1-4/1-1), III (1-3/0-0),
IV (1-3/1-0). Genua I-III (1-4/0-1), IV (1-3/0-0). Tibiae I-II (1-3/2-1),
III-IV (1-3/2-0). Tarsi —17.17.17 (mv absent; av; and pv, stronger, with
opposed points).
_Capitular setae and all hypostomal setae present (Fig. 192). Deutosternum
with about eight denticles in single file. Palpal setal formula 1.2(8).4.8.
Chelicerae bulbous in basal two-fifths, with chelate portion occupying one-
twentieth of total length. Minute tritosternal remnant present.
PTILONYSSUS RHIPIDURAH Domrow
Ptilonyssus rhipidurae Domrow, 1966, Proc. Linn. Soc. N.S.W., 90: 199.
Previous record.—Grey fantail, Rhipidura fuliginosa (Sparrman) (Musci-
capidae, Passeriformes), Esk. Also Innisfail.
PTILONYSSUS MACCLUREI Fain
Ptilonyssus macclurei Fain, 1963, Revue Zool. Bot. afr., 68: 72; Domrow,
1964, Acarologia, 6: 609, 617; 1966, Proc. Linn. Soc. N.S.W., 90: 201.
Previous record.—Willie wagtail, Rhipidura leucophrys (Latham) (Musci-
capidae, Passeriformes), Palen Ck., Brisbane, and Esk. Also Kowanyama.
PTILONYSSUS PSOPHODAE Domrow
Ptilonyssus psophodae Domrow, 1964, Acarologia, 6: 29, 618.
Previous record.—Eastern whipbird, Psophodes olivaceus (Latham)
(Falcunculidae, Passeriformes), Upper Brookfield. Also Esk and Innisfail.
PTILONYSSUS GRALLINAE Domrow
Ptilonyssus grallinae Domrow, 1964, Acarologia, 6: 611, 618.
Previous record.—Magpie-lark, Grallina cyanoleuca (Latham) (Gralli-
nidae, Passeriformes), Brisbane and Condamine. Also Urbenville (N.S.W.),
Esk, Charleville, Chelona and Kowanyama.
PTILONYSSUS RUANDAE Fain
Ptilonyssus ruandae Fain, 1956, Revue Zool. Bot. afr., 538: 395; 1957,
Annls Mus. r. Congo belge Sér. 8v0, 60: 109; Domrow, 1964, Acarologia,
6: 608. Ptilonyssus ruandae alcippei Fain and Nadchatram, 1962, Bull.
Annls Soc. r. ent. Belg., 98: 279.
Previous record.—Grey-backed silvereye, Zosterops lateralis (Latham)
(Zosteropidae, Passeriformes), Brisbane. Also Wilson’s Peak, Mt Jukes,
and Innisfail.
PTILONYSSUS DICAEI Domrow
Ptilonyssus dicaei Domrow, 1966, Proc. Linn. Soc. N.S.W., 90: 201.
Previous record.—Mistletoe-bird, Dicaewm hirundinacewm (Shaw)
(Dicaeidae, Passeriformes), Mt Jukes.
PTILONYSSUS CINNyRIS Zumpt and Till
(Fig. 81)
Ptilonyssus cinnyris Zumpt and Till, 1955, J. ent. Soc. sth. Afr., 18: 78;
Fain, 1957, Annls Mus. r. Congo belge Sér. 8vo, 60: 86; 1958, Bull. Soc. r.
Zool. Anvers 9: 7; Nadchatram, McClure, and Lim, 1964, J. fed. Malay St.
Mus.,9: 104.
ROBERT DOMROW 367
This parasite of African and Malayan sunbirds may now be recorded
from the only Australian member of the family: 3 9 2 and 1 protonymph
from yellow-breasted sunbirds, Cyrtostomus frenatus (Miiller) (Nectariniidae,
Passeriformes), Innisfail, 2.viii and 17.ix.1965, R. D., H. I. MeD., and J. 8. W.
The podonotum is shown in Fig. 81.
PTILONYSSUS MYZANTHAN Domrow
Ptilonyssus myzanthae Domrow, 1964, Acarologia, 6: 603, 617.
_ Previous records (both Meliphagidae, Passeriformes).—Noisy miner,
Myzantha melanocephala (Latham), Condamine. Also Esk. Little wattle-bird,
Anthochaera chrysoptera (Latham), Palen Ck.
New host record.—Yellow-throated miner, M. flavigula Gould, Charleville,
19.i.1967, J. N. and J. S. W. (5 2 2, 1 protonymph) ; Winbin Ck., 20.i1.1966,
Rats Dio, Me candy os. VE (2 2).
PTILONYSSUS PHILEMONI Domrow
(Figs 187-189)
Ptilonyssus philemoni Domrow, 1964, Acarologia, 6:600, 617; 1965,
Tbid., 7: 460.
Previous records (all Meliphagidae, Passeriformes).—Blue-faced honey-
eater, Hntomyzon cyanotis (Latham), Logan Village and Condamine. Also
Esk and Kowanyama. Noisy friar-bird, Philemon corniculatus (Latham),
Logan Village. Also Palen Ck., Esk, Charleville, and Mt Jukes. Little
friar-bird, P. citreogularis (Gould), Brisbane, Samford, and Esk. Also
Charleville, Chelona, Longreach Lagoon, and Kowanyama.
New host records (all Meliphagidae).—Striped honeyeater, Plectorhyncha
lanceolata Gould, Esk, 10.vii.1967, R. D. and B. H. K. (11 2 9, 4 nymphs);
Charleville-Cunnamulla road, 19.v.1965, H. A. S.. D. J. M., and B. H. K.
(82 2). Lesser Lewin honeyeater, Meliphaga notata (Gould), Innisfail, 3 and
22.vi, and 15.ix.1965 (17 2 9,146). Graceful honeyeater, M. gracilis (Gould),
Innisfail, 30.vi, 5—21.viii, and 14.ix.1965, G. J. B., R. D., H. I. MeD., and
SIS. (2c O798)"
The new specimens from Plectorhyncha Gould are quite typical of the
original description, although in one specimen the pygidial shield shows
far to one side a very narrow longitudinal fissure with striate cuticle, and
may fairly be said to be divided.
However, the specimens from Meliphaga Lewin, although they show the
same evenly tapering chelicerae and cordate podonotal shield with 5-6
characteristically arranged pairs of setae figured for P. philemoni Domrow
(1964c), may be separated by the following features: (i) the adanal setae
are set well behind the anus; and (i) the pygidial shield is divided, the
two longitudinal fragments being widely separated. Other minor differences
are detailed in the description below.
Female.—Idiosoma 583-—660n long when unengorged, but up to 825—-990p
when fully fed and mounted; weakly cleft posteriorly. Podonotal shield
(Fig. 189) usually 220-231, long, 170-176» wide, but occasionally smaller
(209-214 long, 165-168. wide); cordate, but margins somewhat eroded,
leaving one or two of three anterolateral pairs of setae free in cuticle; two
discal pairs set in stronger arch than in original series; posterior pair
submarginal. Surface irregularly granulate, with very distinct muscle
insertions. Cuticle between narrowed posterior third of shield and peritremes
with three pairs of setae. Two mesonotal shieldlets accompanied by eight
setae arranged 4.4. Halves of pygidial shield (Fig. 188) elongate, widely
separated, each with terminal seta; preceded by arch of six setae.
368 THE NASAL MITES OF QUEENSLAND BIRDS
Sternal shield somewhat stronger than in original series, with some
granulation, usually bearing SI and II submarginally, and with two pores
between SII; SI not as strong as pygidial setae, SII larger, and SIII
approaching anteroventral opisthosomal setae in size. Metasternal setae
absent. Genital shield typical. Anal shield (Fig. 187) elongate, 140-162p
long, 40-46 wide; anus well forward, and well in front of three subequal
and weak anal setae. Ventral body cuticle with nine to 14 (usually 10-12)
pairs of stout setae.
Setae on coxae and trochanters slightly more expanded than in original
series; those on ventral surface of other segments (including av; and pv;
on tarsi II-IV) somewhat weaker, but still with delicate tips. Setae on
dorsum of legs weaker, evenly tapering. Coxae 2.2.2.1. Trochanters I-II
(1-0/2-1), IIT (2-0/2-0), IV (2-0/4-0). Femur I (2-4/1-2), II (1-4/2-1),
III-IV (1-3/1-0). Genua I-III (1-4/0-1), III (1-4/0-2), IV (1-8/1-0).
Tibiae (1-3/2-1). Tarsi —-.17.17.17 (mv absent). The formulae of the original
series differ in the following respects: trochanter IV with five setae; basifemur
I often with tenth seta ventrally; femur III with four to six setae; genua
ITI-IV usually (1-4/0-1) (1-3/0-0), but (1-4/0-2) (1-38/1-0) in large
specimens from P. corniculatus.
Capitulum as in original series except as follows: basis without spinulae
at ventroexternal angles; hypostomal setae very weak, anteriormost pair
absent. Palpal setal formula 1.2.4.8 in both series.
Male.—Idiosoma 473» long. Dorsum as in female.
Venter essentially as in female, except for genital aperture in front
of SI (area between coxae IV not clear). Anal shield 135 x 40y, flanked by
five stout setae on one side and six on other.
Legs and capitulum undistinguished.
PTILONYSSUS BALIMOENSIS Sakakibara
(Figs 195-203)
Ptilonyssus balimoensis Sakakibara, 1968, J. med. Ent., 5: 17.
T had already figured and described this striking new species before seeing
the above reference, and therefore restrict the present text to additional
descriptive notes and figures. The podonotal shield immediately recalls that
of P. philemoni Domrow (1964c), but the two species may easily be separated
by the shape of the cheliceral shafts, and the setation of the ventral face
of both opisthosoma and palpi.
The new material comprises 2 2 2 from a Macleay honeyeater, Meliphaga
macleayana (Ramsay) (Meliphagidae, Passeriformes), Ella Bay, Innisfail,
12.viii.1965, R. D. and J. S. W.
Female.— Idiosoma (Figs 195-196) 1300» long in slightly compressed
material. Podonotal shield (Fig. 197) broadly cordate, as long as wide;
shagreened over entire surface (except for narrow marginal strip), showing
quite strongly marked muscle insertions, and bearing two subposterior pores
and seven pairs of setae. Pygidial shield (Fig. 199) longitudinally shagreened
(particularly in mid-line), bearing four or five pores and two spinose pygidial
Setae. All dorsal setae minute, particularly on podosoma.
Sternal shield without pores. Genital shield longitudinally shagreened,
with indications of muscle insertions. Anal shield (Fig. 198) shagreened
laterally. Ventral cuticle with 26-31 setae on either side, all of which are
very strong except one midanterior pair.
Coxa IT without anterodorsal process, but anterior seta borne on tubercle.
Setae on ventral aspect of legs considerably longer and stronger than those
ROBERT DOMROW 369
on dorsal, occasionally bifid at extreme tip (Figs 202-208). Coxae 2.2.2.1.
Trochanters I-II (1-0/2-1), III (2—0/2-0), IV (2-0/3-0). Femur I (2-4/3-2)
(including one seta present ventrally on basifemur), II (1-4/2-1), III
(1-4/1-0), IV (1-3/1-0). Genua I-III (1-4/0-1) (but II with only 3 d
once), IV (1-3/0-0). Tibiae I-II (1-3/2-1), III (1-8/2-0) in one specimen
= 7B eee EF > >
-
Figs 195-203. Ptilonyssus balimoensis Sakakibara (? from Meliphaga macleayana) —
195-196, Dorsal and ventral views of idiosoma. 197, Podonotal shield. 198, Anal shield.
199, Pygidial shield. 200-201, Dorsal and ventral views of capitulum. 202-203, Dorsal
and ventral views of leg IV.
370 THE NASAL MITES OF QUEENSLAND BIRDS
(1-3/1 av-0) in other, IV (1-3/2-0) (but with only 1 v, and that av, once).
Tarsi—.17.17.17 (mv absent; avi and pv; stronger on tarsi I-IV). Ambulacrum
I little different from II-IV, all with two subequal claws and diaphanous
pulvillus.
Basis capituli (Figs 200-201) slightly teratological in both specimens,
showing two setae set in contiguous or single alveolus on left side only,
rather than typical 1.1; spinulose externally near trochanteral articulation.
Tritosternum absent, but deutosternal groove present, showing about eleven
uni-| or bidentate denticles. Palpi five-ssegmented, with setal formula
1.3.4(5).8(7) ; four external setae on femur-tibia preternaturally long for
genus, sometimes bifid at extreme tip. Tarsus weak, completely obscured
dorsally by tibia; with about five weak setae and small bifid claw. Chelicerae
suddenly attenuate in distal half; chelate portion occupying one-twentyfifth
of total length.
PTILONYSSUS MYZOMELAE Domrow
Ptilonyssus myzomelae Domrow, 1965, Acarologia, 7: 455.
Previous record.—Scarlet honeyeater, Myzomela sanguinolenta (Latham)
(Meliphagidae, Passeriformes), Samford. Also Esk.
New host record.—Dusky honeyeater, M. obscura Gould, Innisfail, 30.vi,
6.vili, and 30.ix.1965, G. J. B., R. D., H. I. MeD., and J. S. W. (3 2 9,1
deutonymph) ; Jordan Ck., 18.viii.1965, R. D. (22 2, 1 protonymph).
Specimens from the type host, M. sanguinolenta, show the pygidial shield
completely lacking, although the short, stumpy pygidial setae (and their
accompanying pores) are easily made out among the elongate, tapering
opisthosomal series. Specimens from M. obscura, however, show widely
separated, small, rounded pygidial shields, each bearing a seta and pore,
although on one side of one specimen the pygidial seta is separated from its
shield and pore by five complete cuticular annulations.
PTILONYSSUS GLICIPHILAE Domrow
(Figs 171-173)
Ptilonyssus gliciphilae Domrow, 1966, Proc. Linn. Soc. N.S.W., 90: 204.
Previous record.—Brown honeyeater, Gliciphila indistincta (Vigors and
Horsfield) (Meliphagidae, Passeriformes), Esk and Chelona. Also Charleville
and Kowanyama.
New host record.—Banded honeyeater, Myzomela pectoralis Gould
(Meliphagidae), Kowanyama, 16, 18, and 23.x.1965, R. D. (11 2 2,12).
The podonotal shield of the new material is very similar to that of the
original. However, apart from host-specificity, the new form is readily
separable by showing the pygidial shield distinctly divided, and two setae
ventrally on basifemur I. Further minor differences are noted in the
description below.
Female.—Idiosoma 780p long in one unengorged specimen, but elongate
and reaching 1320-1430» in fully fed, mounted material; terminal bifurcation
quite evident in life, even to naked eye. Podonotal shield (Fig. 173) usually
290-308 long and 187-192» wide, but extreme range 286-319. long and
176-205 wide; more parallel-sided in posterior half, and with anteriormost
pair of median setae set further back than in original series; surface strongly
granulate, with well-defined muscle insertions. Arrangement of adjacent
Setae and shieldlets around podonotal shield and on mesonotum as in original
series. Halves of pygidial shield (Fig. 171) small, well separated, each with
Seta and pore; flanked by three or four pairs of stronger setae.
ROBDRT DOMROW 371
Venter as in original series, but anal shield (Fig. 172) narrower (220-2538,
long, 52-554 wide) and less well-defined anteriorly. Genital shield on raised
lobe between coxae IV in engorged specimens.
Coxae 2.2.2.1. Trochanters I-II (1-0/2-1), III (2-0/2-0), IV (2-1/2-0).
Femur I (2-4/3-2) (including two setae ventrally on basifemur), IT (1-4/2-1),
III (1-3/2-0), IV (1-38/1-0). Genu I (1-4/0-1), II (1-4/1-1), III (1-4/1-2),
IV (1-8/1-0). Tibiae (1-3/2-1). Tarsi -.17.17.17 (mv absent; av, and pri
as in original series). The original series shows a similar chaetotaxy, except
femur I (1-4/1-2) (ie. no setae ventrally on basifemur), III (1-3/1-0) ;
genu IIT (1-4/1-1). Setae on ventral face of legs similar to those on coxae,
but those on dorsum rather weaker. Ambulacrum I only very slightly weaker
than II-I[V. Coxa IT with strong anterodorsal process.
Capitulum as in original series, but setal formula of palpi 1.3(2).4.9
due to presence of additional seta dorsally on femur and tibia. Tritosternum
absent.
Male.—Idiosoma 596p long, evenly rounded posteriorly. Dorsum entirely
as in female, but podonotal shield smaller, 219 x 120p.
Sternal shield as in female, but extended into diffuse genital area between
coxae IV; genital setae free in cuticle. Sexual aperture in front of SI. Anal
shield as in female, but only 120» long; surrounded by five pairs of setae
rather than 10-12 pairs in female.
Leg setation as in female, but with five setae on one trochanter II; av,
and pv; on tarsi II-IV sharply pointed, though still upturned at tip.
Capitulum as in female. Chelicerae not attenuate, spermatodactyl
occupying two-fifths of total length of 120».
PTILONYSSUS LYMOZEMAR Domrow
Ptilonyssus lymozemae Domrow, 1965, Acarologia, 7: 453.
Previous record.—Scarlet honeyeater, Myzomela sanguinolenta (Latham)
(Meliphagidae, Passeriformes), Logan Village and Samford. Also Esk.
PTILONYSSUS STOMIOPERAE Domrow
Ptilonyssus stomioperae Domrow, 1966, Proc. Linn. Soc. N.S.W., 90: 206.
Previous records (both Meliphagidae, Passeriformes ).—Yellow honeyeater,
Meliphaga flava (Gould), Kowanyama. White-gaped honeyeater, Stomiopera
unicolor (Gould) Kowanyama.
PTILONYSSUS THYMANZAE Domrow
Ptilonyssus thymanzae Domrow, 1964, Acarologia, 6: 604, 617; 1966,
Proc. Linn. Soc. N.S.W., 90: 204.
Previous records (all Meliphagidae, Passeriformes).—Lewin honeyeater,
Meliphaga lewinii Swainson, Esk. Also Wilson’s Peak and Innisfail. Lesser
Lewin honeyeater, M. notata (Gould), Innisfail. Yellow-faced honeyeater,
M. chrysops (Latham), Samford. Noisy miner, Myzantha melanocephala
(Latham), Logan Village and Condamine. Also Brisbane and Esk. Little
wattle-bird, Anthochaera chrysoptera (Latham), Palen Creek.
New host records (all Meliphagidae).—White-plumed honeyeater, Meli-
phaga penicillata Gould, Charleville, 10 and 24.1.1967, R. D., J. N., and
J.S. W. (42 2); Winbin Ck., 21.1.1966, R. D., D. J. M., and J. 8. W. (22 2).
Yellow-throated miner, Myzantha flavigula Gould, Winbin Ck., 20.1.1966, R. D..
D. J. M., and J. S. W. (2 2 2). Spiny-cheeked honeyeater, Acanthagenys
rufogularis Gould, Charleville, 14.1.1966 and 1.ii.1967, R. D., D. J. M., and
J. Ss Waa) O42);
372 THE NASAL MITES OF QUEENSLAND BIRDS
PTILONYSSUS MELIPHAGAE Domrow
Ptilonyssus meliphagae Domrow, 1964, Acarologia, 6: 606.
Previous record.—Yellow-faced honeyeater, Meliphaga chrysops (Latham)
(Meliphagidae, Passeriformes), Mt Cotton. Also Hsk.
New host record.—White-throated honeyeater, Melithreptus albogularis
Gould (Meliphagidae), Esk, 6.x.1966 and 17.iv.1967, R. D., B. H. K., and
J.S. W. (4 2 2); Kowanyama, 15.iii.1966, R. D. (12 ).
The pygidial complex of the original specimen from Meliphaga was not
detected, but I have since seen two more females from this host. These show
the pygidial shield widely divided into two elongate fragments little wider
than the alveoli of the pygidial setae. A pore accompanies each seta.
The specimens from Melithreptus Vieillot all show an entire, transverse
pygidial shield, although in two specimens it is narrowly and asymmetrically
divided as in the specimen of P. philemoni discussed above. Indeed, in one
specimen, the teratology affects the entire distal portion of the opisthosoma,
as ventrally the anal shield is also atypical, lacking the postanal seta.
PTILONYSSUS NUDUS Berlese and Trouessart
Ptilonyssus nudus Berlese and Trouessart, 1889, Bull. Biblioth. scient.
Ouest, 2: 180; Berlese, 1892, “Acari, myriapoda, et scorpiones hucusque in
Italia reperta, ordo Mesostigmata’”’ (Patavii), fasc. 58, No. 10 (2); de
Castro and Pereira, 1947, Archos Inst. biol., S Paulo, 18: 127; Bregetova,
1951, Parazit. Sb., 13: 118; Porter and Strandtmann, 1952, Tex. J. Sci., 4: 394;
George, 1961, J. Kans. ent. Soc., 834: 116; Fain, 1963, Bull. Amnls Soc. ae ent.
Belg., 99: 171; nec Hirst, 1916, a Zool. Res., 1:78. Ptilonyssus (Ptilonyssus)
nudus, de Castro, 1948, Archos Inst. biol., S Paulo, 18 : 260.
Previous records (both Fringillidae, Passeriformes) —Tree-sparrow (intro-
duced), Passer montanus (Linnaeus). House-sparrow (introduced), P.
domesticus (Linnaeus).
I have been able to examine specimens of this species from Europe and
the United States.
PTILONYSSUS STURNOPASTORIS Fain
(Figs 204-213)
Ptilonyssus sturnopastoris Fain, 1963, Acarologia, 5:1.
This species, originally described from an Indian starling (Sturnopastor
Blyth), may now be recorded as follows: 2 29 9 and 2 protonymphs from
Australian shining starlings, Aplonis metallica (Temminck) (Sturnidae,
Passeriformes), Ella Bay, 4.viii.1965, R. D. and J. S. W.
PTILONYSSUS SPHECOTHERIS Domrow
Ptilonyssus sphecotheris Domrow, 1964, Acarologia, 6: 613.
Previous record.—Southern figbird, Sphecotheres vieillotii Vigors and
Horsfield (Oriolidae, Passeriformes), Brisbane. Also Samford.
New host records.—Yellow figbird, S. flaviventris Gould, Ella Bay,
4.vili.1965, R. D. and J. S. W. (41 2 2,6 ¢ 6,1 deuto-, and 1 protonymph).
Great bower-bird, Chlamydera nuchalis (Jardine and Selby) (Ptilonorhyn-
chidae, Passeriformes), Kowanyama 18.iii.1966, R. D. (192).
I have not been able to distinguish the lattermost specimen from paratypes
of this species. Its presence in a ptilonorhynchid rather than the usual
oriolid is not due to any error of labelling, either in the field or the laboratory.
As with the single specimens above of P. cractici from Dacelo and R. rhino-
lethrum from Tringa, I was quite aware at the time of collection that a novelty
had been obtained.
ROBERT DOMROW 373
PTILONYSSUS TROUDSSARTI (Hirst)
Rhinonyssoides trouessarti Hirst, 1921, Proc. zool. Soc. Lond., 1921:
770. Ptilonyssus trouessarti, Fain and Hyland, 1962, Ann, Mag. nat. IHist.,
(13) 5: 346; Domrow, 1964, Acarologia, 6: 608. Ptilonyssus orioli Fain, 1956,
Revue Zool. Bot. afr., 53: 144; 1957, Annls Mus. r. Congo belge Sér. 8vo, 60:
114; 1962, Bull. Annls Soe. r. ent. Belg., 98: 264; Strandtmann, 1960, J. Kans.
ent. Soc., 88: 150.
ht Ow
Figs 204-213. Ptilonyssus sturnopastoris Fain (2 from Aplonis metallica) —204-205,
Dorsal and ventral views of idiosoma. 206, Podonotal shield. 207-208, Variations in
Pygidial shield. 209, Anal shield. 210, Ventral view of capitulum (left palp in dorsal
F view). 211, Chelicera. 212-218, Dorsal and ventral views of leg III.
374 THE NASAL MITES OF QUEENSLAND BIRDS
Previous records (both Oriolidae, Passeriformes).—Olive-backed oriole,
Oriolus sagittatus (Latham), Logan Village. Also Esk and Innisfail. Yellow
oriole, O. flavocinctus (King), Kowanyama.
Domrow (1946c) has discussed the misidentification of the original host
of this species.
PTILONYSSUS NOVAEGUINEAE (Hirst)
Rhinonyssus novae-guineae Hirst, 1921, Proc. zool. Soc. Lond., 1921:
769. The original spelling (with hyphen) is corrected above under Art. 32.
Passeronyssus novae-guineae, Fain, 1960, Revue Zool. Bot. afr., 61: 318.
Ptilonyssus novae-guineae, Fain and Hyland, 1962, Ann. Mag. nat. Hist., (13)
5: 346.
Previous record.—Magnificent rifie-bird, Ptiloris magnificus (Vieillot)
(Paradiseidae, Passeriformes).
PTILONYSSUS CERCHNEIS Fain
Ptilonyssus cerchneis Fain, 1957, Annls Mus. r. Congo belge Sér. 8vo,
60: 133; Strandtmann, 1962, Proc. ent. Soc. Wash., 64: 100; Bregetova,
1964, “Some problems of evolution of the rhinonyssid mites” (Nauka:
Leningrad), p.5; Domrow, 1965, Acarologia, 7: 457.
Previous records.—Brown hawk, Falco berigora Vigors and Horsfield
(Falconidae, Falconiformes), Esk. Also Kowanyama. Nankeen kestrel, F.
cenchroides Vigors and Horsfield, Mt Jukes. Also Esk.
Most of my specimens were heavily engorged when collected, and the
opisthosomal cuticle has accordingly ruptured and crumpled during mounting
procedures. One unfed specimen, however, does show the two areas of
non-striate cuticle on the opisthonotum described by Strandtmann (1962),
as well as a second, smaller pair slightly behind, and outside the first pair.
Neither pair bears a seta and associated pore, and it would therefore seem
that Strandtmann is correct in questioning whether these structures belong
to the pygidial complex. I have examined 22 @from the United States.
Genus Sternostoma Berlese and Trouessart
Sternostoma Berlese and Trouessart, 1889, Bull. Biblioth. scient. Ouest,
2: 126; Furman, 1957, Hilgardia, 26: 474. Type-species Sternostoma
cryptorhynchum Berlese and Trouessart, 1889, Loc. cit., 127. Sternostomum
Trouessart, 1895, Rev. Sci. nat. appl., 42: 392. Unjustified emendation under
Art. 33. Furman (1957) has pointed out the error of Vitzthum (1935) and
his school in considering Sternostoma and Sternostomum distinct genera, even
though Trouessart had explicitly stated that the type-species of the former
was also that of his unjustified emendation. Nor is either name to be
confused with Rhinonyssus Trouessart, 1894, v. supra, type-species Rhinonyssus
coniventris Trouessart, 1894. Agapornyssus Gretillat, Capron, and Brygoo.
1959, Acarologia, 1: 376. Type-species Agapornyssus faint Gretillat et al.,
1959, Loc. cit., 876. Sternostomoides Bregetova, 1964, “Some problems of
evolution of the rhinonyssid mites” (Nauka: Leningrad). Sternostomoides
is not mentioned in the text of this article, but occurs in an accompanying
photograph of a table on host-specificity. This does not constitute publication
under Art. 8; and, in any case, as no type-species is given, Sternostomoides
is unavailable here under Art. 18. Also 1965, Hnt. Obozr., 44: 709. Type-species
Sternostomum technawi Vitzthum, 1935, J. Orn., Lpz., 83: 569. New synonymy.
Rhinosterna Fain, 1964, Revue Zool. Bot. afr., 70: 125. Type-species
Rhinosterna aymarae Fain, 1964, Loc. cit., 126. New synonymy. Sternoecius
Fain and Aitken, 1967, Bull. Inst. r. Sci. nat. Belg., 43: 24. Type-species
Sternoecius piprae Fain and Aitken, 1967, Loc. cit., 24. New synonymy.
ROBERT DOMROW 375
Sternostomoides was erected for a group of three parasites of turdids (all
considered conspecific below), which lack the opisthonotal shield. However,
the existence of S. inflatwm Fain (1968e) and S. duwreni Fain (1956, 1957e),
also from turdids, in which the opisthonotal shield is eroded and complete,
respectively, makes the synonymy clear.
The cheliceral modifications in the last two taxa should be considered
only of specific value, since the two species involved are otherwise typical
of Sternostoma.
Key to females of Australian species of STERNOSTOMA
il. Podonotal shield evenly rounded anteriorly. Opisthonotal shield at most
accompanied by small mesonotal shieldlets, or absent .............. 2
Podonotal shield deeply excavated anteriorly. Opisthonotal shield accompanied
by two extensive lateral shields ..,............... fulicae Fain and Bafort
Bl) ree OMIStHOnOtale SHICIG: PRESEN ono srecteeileris syecccucs s onic cleus s cbetaus has Mefaraieta ates 3
Opischonotaleshiclawansenitiee smi-rere clerciets lstevale wieretcts oo) otalel ote technaui (Vitzthum)
3 (2). Posterolateral margin of podonotal shield with four setae (two medial and
one in each corner) whee Nee oiceSeNS SYD ApS CPETSR SY PRESS ohosie. Saale ole tie BEANE oe Aes 4
: Posterolateral margin of podonotal shield with two setae (medial) ...... 9
4 (3). Two mesonotal shieldlets present. Palpal tibiotarsus twice as long as wide
EOI CQOIS- Claro CIO DO C ORROIDAE: cicIRoScibur MEIC CD CAO CRO OIC ROO De thienponti Fain
Mesonotal shieldlets absent. Palpal tibiotarsus at most only slightly longer
CHAT WADE fart peers hacis chase sea Gide Ae ops = cee: = thee eal esas Aistdeie eb sche. 5
5 (4). Setae dly-., GVy-2, PVi-2, and pl,-. on tarsi II-IV variously formed, but not as
HOMON TS). poe ooo ooUr EVV A PALE. ace Rue Tide: TALESPEN ccs. Sete fadaie care folete na: alt tertioie 6
Setae a@ly-., @Vys, PUi-2, and pl.-. on tarsi II-IV slender, with expanded tips
SURNON AS et Nao picket becerran oust alan ace-ahe Prato von her cu etes cel chicka: aka eae Kon seks ct Lavtote\a en cheite cooremani Fain
6 (5). Onigihonotat shield subquadrate. Setae Ql,-1, GUVy-.2, DVy-2, and pl,-, on tarsi
INI Qe Mehicahtey rey PS eels rea, iS key ee eta BR Biota Mey CS RCE cuculorum Fain
Opisthonotal shield at least one and a half times as long as wide Setae
Glo, AVy2, PViz, and pl-, on tarsi II-IV variously formed, but not as
BDOVERE SE ae hace ae Ree edna tele s Sete PA Re SRAEE . Tbe oe a EE. shale qf
7 (6). Dorsal shield heavily and evenly punctate. Palpal tibial sensory rods minute
BP eho oo Oe Ono UB cho Oc G05 ct TERE. Seton 1G DC cha RIOTS BiBie one Reece er eee or ae a 8
Dorsal shield with strong subhexagonal reticulation giving the effect of
honeycomb. Palpal tibial sensory rods more than half as long as
GUDLOCATSUS LPS Be eR eS TS eS Gre gliciphilae Domrow
8 (7). Sternal shield weakly and evenly granulate. Some of setae Ql, AVy-2, DV1-s,
and pl-, on tarsi II-IV peculiarly crimped .................. dureni Fain
Sternal shield with distinct transverse reticulation. Setae dl-., @V,;-2, DUi-2, and
pl,-. on tarsi II-IV minute, unmodified .......... tracheacolum. Lawrence
9 (3). Lateral margins of podonotal shield evenly convex ................20005- 10
Lateral margins of podontal shield triconcave ................ paddae Fain
10 (9). Podonotal shield with five pairs of setae. Mesonotal shields absent ...... ill
Podonotal shield with four pairs of setae. Mesonotal shields present........
PTR Bane Che PeT eT a eT Nee es, RE NGKE. SS CL oie Rie Te OhGke welele REN Bie eS elel eel shone voydi Strandtmann
11 (10). With one pair of setae between genital and anal shields. Setae al.-., GVy-2, DUro,
and pi,-, on tarsi Il-IV wnmodified -................... neositiae, D. sp.
With two pairs of setae between genital and anal shields. Setae al-,, avj-.,
fivin, Enel Wee, Cit Ven EIN Groene 5 e9e od BsbssSebkaeeadsusgeuccuede
BUS A Oe PARES. 5 aks alld cryptorhynchum Berlese and Trouessart
STERNOSTOMA THIENPONTI Fain
Sternostoma thienponti Fain, 1956, Revue Zool. Bot. afr., 53: 152; 1957,
Annls Mus. r. Congo belge Sér. 8vo, 60: 68; Domrow, 1965, Acarologia, 7: 449;
1966, Proc. Linn. Soc. N.S.W., 90:194. Sternostoma thieponti (sic)
Strandtmann, 1960, J. Kans. ent. Soc., 33: 135.
Previous records (all Passeriformes).—Black-backed magpie, Gymnorhina
tibicen (Latham) (Cracticidae), Condamine. Black butcher-bird (black and
red phases), Cracticus quoyi (Lesson and Garnot), Innisfail. Spangled drongo,
Chibia bracteata (Gould) (Dicruridae).
New host record.—Pied butcher-bird, Cracticus nigrogularis (Gould),
-Kowanyama, 8.iv.1965, R. D.and J.S. W. (7 2 @).
376 THE NASAL MITES OF QUEENSLAND BIRDS
STERNOSTOMA COOREMANI Fain
Sternostoma cooremani Fain, 1956, Revue Zool. Bot. afr., 53: 154; 1957,
Annls Mus. r. Congo belge Sér. 8vo, 60: 72; Domrow, 1965, Acarologia, 7:
448; 1966, Proc. Linn. Soc. N.S.W., 90: 192. Sternostoma cooremani halcyom
Fain and Nadchatram, 1962, Bull. Anitls Soc. r. ent. Belg., 98: 280.
Previous records (all Coraciiformes).—Laughing kookaburra, Dacelo
gigas (Boddaert) (Alcedinidae), Condamine. Also Esk. Mangrove-kingfisher,
Halcyon chloris (Boddaert) (Alcedinidae). Rainbow-bird, Merops ornatus
Latham (Meropidae), Esk. Also Windorah.
STERNOSTOMA CUCULORUM Fain
Sternostoma cuculorum Fain, 1956, Revue Zool. Bot. afr., 53: 155; 1957,
Annls Mus. r. Congo belge Sér. 8vo, 60: 74; Nadchatram, McClure, and
Lim, 1964, J. fed. Malay St. Mus., 9: 106; Domrow, 1965, Acarologia, 7: 449.
Sternostoma cuculorum var. urolestis Fain, 1959, J. ent. Soc. sth. Afr.,
22: 32. See Art. 45. New synonymy. Sternostoma laniorum Fain, 1956,
Revue Zool. Bot. afr., 53: 156; 1957, Annls Mus. r. Congo belge Sér. 8vo,
60: 76; Domrow, 1965, Acarologia, T: 449; 1966, Proc. Linn. Soc. N.S.W.,
90: 193. New synonymy. Sternostoma laniorum var. batis Fain, 1957, Annls
Mus. r. Congo belge Sér. 8vo, 60: 77; Fain and Nadchatram, 1962, Bull. Annls
Soc. r. ent. Belg., 98: 281; Strandtmann, 1960, J. Kans. ent. Soc., 33: 185.
See Art. 45. New synonymy. Sternostoma batis, Fain, 1962, Revue Zool.
Bot. afr., 66: 146; Nadchatram, McClure, and Lim, 1964, J. fed. Malay St.
Mus., 9: 106. Sternostoma zosteropus Domrow, 1966, Proc. Linn. Soc. N.S.W.,
90: 194. New synonymy.
Until more tangible characters can be found to divide the above taxa,
I would prefer to consider them conspecific. The character of the claws
originally noted for S. zosteropus does not, in fact, hold good. It may be
that in this (and in one other case, viz. Ptilonyssus motacillae, v. supra) ;
I have been too radical in my synonymy, but it is virtually impossible to
key their component taxa as presently described.
Previous records.—Fan-tailed cuckoo, Cacomantis pyrrhophanus (Vieillot)
(Cuculidae, Cuculiformes), Esk. Also Innisfail and Kowanyama. Willie
wagtail, Rhipidura leucophrys (Latham) (Muscicapidae, Passeriformes), Esk.
Also Kowanyama. Leaden flycatcher, Myiagra rubecula (Latham) Musci
capidae), Innisfail. Pale-yellow robin, Hopsaltria capito Gould (Musci-
capidae), Innisfail. Rufous strike-thrush, Colluricincla megarhyncha (Quoy
and Gaimard) (Pachycephalidae, Passeriformes), Innisfail. Crested bellbird,
Oreoica gutturalis (Vigors and Horsfield) (Falcunculidae, Passeriformes),
Mitchell. Grey-backed silvereye, Zosterops lateralis (Latham) (Zosteropidae,
Passeriformes), Mt. Jukes.
New host records—Brush cuckoo, C. variolosus (Vigors and Horsfield),
Kowanyama, 14.iv.1965, R. D. (2229). Koel, Hudynamys orientalis
(Linnaeus) (Cuculidae), Esk, 6.x.1966, R. D. and J. 8. W. (142 2). Australian
pied flycatcher, Arses kawpi Gould (Muscicapidae), Innisfail, 5.viii.1965, R. D.
and J.S. W. (19 @ 2). Spectacled flycatcher, Monarcha trivirgata (Temminck)
(Muscicapidae), Innisfail, 31.viii.1965, H. I. McD. (10 2 @).
STERNOSTOMA DURENI Fain
Sternostoma dureni Fain, 1956, Rev. Zool. Bot. afr., 53: 153; 1957, Annls
Mus. r. Congo belge Sér. 8vo, 60: 68.
This species, whose type-host is an African thrush, may now be recorded
from Australia: 42 2 from an Australian ground-thrush, Oreocincla lunulata
ae) (Turdidae, Passeriformes), Wilson’s Peak, 15.v.1967, R. D. and
x EL. Ke
ROBHRT DOMROW 377
STHRNOSTOMA TRACHHPACOLUM Lawrence
Sternostoma tracheacolum Lawrence, 1948, J. Parasit., 34: 366;
Bregetova, 1951, Parazit. Sb., 18: 116; Furman, 1957, Hilgardia, 26: 478;
Fain, 1957, Annls Mus. r. Congo belge Sér. 8vo, 60: 65; Fain and Hyland,
1962, Parasitology, 52: 404; Strandtmann, 1960, J. Kans. ent. Soc., 33: 137;
Domrow, 1965, Acarologia, 7: 449; 1966, Proc. Linn. Soc. N.S.W., 90: 194;
Murray, 1966, Aust. vet. J., 42: 262. Sternostoma medda Lombardini, 1953,
Redia, 38: 187. Sternostoma castroae Fain, 1956, Revue Zool. Bot. afr.,
58: 398. Agapornyssus fain Gretillat, Capron, and Brygoo, 1959, Acarologia,
Le oG:
Previous records (all Passeriformes except the first).—Budgerygah
(aviary-bred), Melopsittacus undulatus (Shaw) (Psittacidae, Psittaciformes).
Common swallow, Hirundo rustica Linnaeus (Hirundinidae). Australian
reed-warbler, Acrocephalus australis (Gould) (Sylviidae). Yellow-breasted
sunbird, Cyrtostomus frenatus (Muller) (Nectariniidae). Goldfinch (intro-
duced), Carduelis carduelis (Linnaeus) (Fringillidae). House-sparrow (intro-
duced), Passer domesticus (Linneaus) (Fringillidae). Canary (introduced
cage-bird), Serinus canaria (Linneaus) (Fringillidae). Gouldian finch (aviary-
bred), Poephila gouldiae (Gould) (Ploceidae), Sydney.
This mite is the cause of severe respiratory complications in cage-birds
(Fain and Carpentier, 1958; Fain and Hyland, 1962a; Murray, 1966; Mathey,
1967).
STERNOSTOMA GLICIPHILAE Domrow
Sternostoma gliciphilae Domrow, 1966, Proc. Linn. Soc. N.S.W., 90: 192.
Previous record.—Brown honeyeater, Gliciphila indistincta (Vigors and
Horsfield) (Meliphagidae, Passeriformes), Esk.
STERNOSTOMA NEOSITTAE, 0D. Sp.
(Figs. 188, 214-218)
Sternostoma sp. Domrow, 1967, Proc. Linn. Soc. N.S.W., 91: 217.
Diagnosis—The new species, which lacks setae in the posterolateral
angles of the podonotal shield, and does not show the digitate, or even
apically expanded setae aly-2, avi1-2, pUi-2, and ply-2 on tarsi II-IV character-
istic of such species as the type, S. cryptorhynchum (well figured from the
original material by Furman, 1957, and Fain, 1957e), may best be compared
with S. boydi Strandtmann (1951) and S. inflatum Fain (1963e). The former
(originally described from the sanderling, Scolopacidae, Charadriiformes)
is Separable by the presence of only four pairs of setae on the podonotal
shield (v. infra), the presence of two pairs of setae, not one, behind the
genital shield, and its relatively unmodified anal shield. The latter (from
a thrush, Turdidae, Passeriformes) is separable by the eroded margins of
its opisthonotal shield, and the presence, again, of two pairs of setae behind
the genital shield.
Types.—Holotype female, sixteen paratype females, and two deutonymphs
enclosing developing males from a striated sittella, Neositta striata (Gould)
(Sittidae, Passeriformes), Kowanyama, 7.iv.1965, R. D. and J. S. W. Holotype
N. I. C.; paratypes R. D., A. F., and R. W. S.
Female—tIdiosoma 4954 long in unengorged specimen, and from 640—
690 in engorged mounted specimens like that illustrated. Podonotal shield
(Fig. 214) 260-270 long, 160-165. wide; anterior quarter narrower and
strongly convex, sides subparallel, and posterior margin rectilinear. Surface
with punctae heavier towards midline, particularly where two rows of muscle
insertions converge. Setae in three anterolateral, and two posteromedian
G
378 THH NASAL MITES OF QUEENSLAND BIRDS
pairs; two pairs of pores present. Opisthonotal shield rectangular, 140-150,
long, 110-115 wide, bearing four setae, and again showing punctae heaviest
near muscle insertions. Stigmata without peritremes, unarmed. Dorsal
cuticle without additional setae. .
216 ew
Figs 214-218. Sternostoma neosittae, n. sp. (Q from Neositta striata) —214-215, Dorsal
and ventral views of idiosoma. 216-217, Ventral and dorsal views of tarsus III. 218,
Ventral view of capitulum (right palp in dorsal view). :
Figs 219-222. Ptilonyssus sittae Fain (protonymph from Climacteris picumnus).—Dorsal
and ventral views of femora-tarsi III-IV.
ROBDRL DOMROW 379
Sternal shield (Fig. 215) longer than wide, somewhat expanded between
coxae If and IIT; surface lightly shagreened; SI-III present in longitudinal
rows, but metasternal setae absent. Genital shield drop-shaped, with distinct
recticulation and punctation; rayed operculum present; genital seta-pore
complex reduced to remnant just off shield. Anal shield terminal, very weakly
sclerotized, and without cribrum. Adanal setae present, but postanal absent.
Ventral cuticle with one pair of setae behind genital shield. All body setae
minute.
Leg setae again minute, barely longer than diameter of ‘their alveoli.
Coxal formula 2.2.2.0. Tarsus I with dorsodistal sensory islet as in Fig.
188. Tarsal formula—.18.18.18 (mv present). Tarsi and ambulacra I-IV
stouter than I (Figs 216-217).
Capitulum undistinguished (Fig. 218). Capitular setae well forward,
with one, or even both of pair frequently absent. HI not detected, II-III
very weak. Dorsodistal rods on tibia very short. Tarsus obsolescent, with
no trace of claw. Tritosternum absent.
STERNOSTOMA CRYPTORHYNCHUM Berlese and Trouessart
Sternostoma cryptorhynchum Berlese and Trouessart, 1889, Bull. Biblioth.
scient. Ouest, 2:127; Furman, 1957, Hilgardia, 26: 475; Fain, 1957, Annls
Mus. r. Congo belge Sér 8vo, 60: 70.
Previous record.—House-sparrow (introduced), Passer domesticus
(Linnaeus) (Fringillidae, Passeriformes).
STRRNOSTOMA PADDAE Fain
Sternostoma paddae Fain, 1958, Bull. Soc. r. Zool. Anvers, 9: 8.
Previous record.—Java sparrow (introduced cage-bird), Padda oryzivora
(Linnaeus) (Ploceidae, Passeriformes).
Fain and Bafort (1963) report this mite associated with severe conjunc
tivitis in this host.
STERNOSTOMA ®oyDI Strandtmann
Sternostoma boydi (sic) Strandtmann, 1951, J. Parasit., 37: 138 (a
solecism now allowable under Rec. 31--the species was named for Dr.
Elizabeth M. Boyd); Furman, 1957, Hilgardia, 26: 480; Fain, 1956, Revue
Zool. Bot. afr., 53: 151; 1960, Tbid., 62: 95; Mitchell, 1961, SWest Nat.,
6: 103.
Previous records (all Charadriiformes).—White-winged black tern,
Chlidonias leucoptera (Temminck) (Laridae). Sooty tern, Sterna fuscata
Linnaeus (Laridae). This record is based on host-relationship (Amerson,
1967), and IT have been unable to obtain specimens to confirm it. Turnstone,
Arenaria interpres (Linnaeus) (Charadriidae). Common sandpiper, Tringa
hypoleuca Linnaeus (Scolopacidae). Wood-sandpiper, 7. glareola Linnaeus.
Greenshank, 7. nebularia (Gunnerus). Sanderling, Crocethia alba (Vroeg)
(Scolopacidae). Knot, Calidris canutus (Linnaeus) (Scolopacidae).
I have examined eight @ of this species from Rwanda and the United
States, all with the podonotal shield showing three pairs of anterolateral
setae in addition to the single posteromedian pair originally figured.
STERNOSTOMA TECHNAUI (Vitzthum)
Sternostomum technaui Vitzthum, 1935, J. Orn., Lpz., 83: 569.
Sternostoma technaui, Furman, 1957, Hilgardia, 26: 482; Fain, 1963, Bull.
Annls Soc. r. ent. Belg., 99:174. Sternostomoides technaui, Bregetova, 1965,
Ent. Obozr., 44: 709. Sternostoma turdi Zampt and Till, 1955, J. ent. Soc. sth.
380 THE NASAL MITES OF QUEENSLAND BIRDS
Afr., 18: 85; Furman, 1957, Hilgardia, 26: 481; Fain, 1962, Bull. Annls Soe. r.
ent. Belg., 98: 264; 1963, [bid., 99: 176. New synonymy. Sternostomoides turdi,
Bregetova, 1965, Hnt. Obozr., 44: 712. Sternostoma spatulatum Furman,
1957, Hilgardia, 26: 480. New synonymy.
Previous records (all Turdidae, Passeriformes).—Blackbird (introduced),
Turdus merula Linnaeus. Song-thrush (introduced), 7. philomelos Brehm.
Australian ground-thrush, Oreocincla lunulata (Latham).
New Australian record.—O. lunulata, Wilson’s Peak, 15.v.1967, R. D.
andy Bt HK 41 79%)2
I do not believe the three nominal species above reflect more than the
intraspecific variation to be expected in any cosmopolitan parasite. All are
parasites of turdids.
STERNOSTOMA FULICAE Fain and Bafort
Sternostoma fulicae Fain and Bafort, 1963, Bull. Annls. Soc. r. ent.
Belg., 99: 474.
Previous record.—Coot, Fulica atra Linnaeus (Rallidae, Gruiformes).
Family EREYNETIDAE
Subfamily SPELEOGNATHINAE
As the type-genus, Speleognathus Womersley (1936), was consistently
so spelled by the original author, his rendering of the family-group name
as “Spelaeognathidae” contravenes Art. 29. An obvious lapsus calami, it
should be corrected under Art. 82.
Comments on the labile nature of most, if not all characters used in
the three most recent classifications of the Speleognathinae (Fain, 19586,
1962d, 1963c) continue to appear (Clark, 1964, 1967; Domrow, 1961, 1965a),
and I therefore accept only the genera of the third of Fain’s systems.
Key to adults of Australian genera of SPELEOGNATHINAE
ils All body and leg setae barbulate. Dorsal shield absent. Eyes normally absent,
but present in B. myzomelae, n. sp. Sensillae weakly clavate. Palpi with
THES MSeSMENTS < oc sciessre ec decueue, ts sve search eatreeveeaopeeeecene Boydaia Womersley
Body and leg setation with an admixture of filamentous setae. Dorsal shield
and eyes absent, weak, or substantial. Sensillae normally filamentous,
but globose in Neoboydaia merops (Fain). Palpi with one, two, or three
SCEMENCS. > ok Gites Ho Be alee ee ere eas asics ade Beles OR Soe eee seer Mg weet
2 (1). Dorsal shield absent or minute. Palpi with one, two, or three segments .... 3
Dorsal shield substantial. Palpi with three segments ......................
Bile ER SR «ea ine Nog AORN COE hd ok CRRA RAE Ee ee Speleognathopsis Cooreman
3 (2). Eyes absent. Palpi with one or two segments ............ Neoboydaia Fain
Eyes present. Palpi with two or three segments .. Ophthalmognathus Dubinin
Genus Boypa1a Womersley
Boydaia Womersley, 1958, Trans. R. Soc. S. Aust., 76: 82. Type-species
Speleognathus sturni Boyd, 1948, Proc. ent. Soc. Wash., 50: 9, nec Boydaa
-angelae Womersley (1953) as stated by Ford (1959) and Brooks and Strandt-
mann (1960). The latter is a species of Lawrencarus Fain (1957c)
(Ereynetidae, Lawrencarinae), a genus inhabiting the nasal passages of
frogs and toads.
Key to adults of Australian species of BoYDAIA
il, Coxal@setae) not tlt Ey postomaly setae 25200. cis telerail teeta alien 2
Coxall setae mal Meals ery postomalmsetaew = Liaraerin einen falconis Fain
21) aloxternal setasonncoxa la normealign arrancones Felder s Gahan 3
Hxternal seta, onecoxay ee mIniten aoe een eerie hirundoae Fain
ROBDRT DOMROW 381
8 (2). Trochanteral setae 1.1.0.0. Femoral setae 6.4.3.2 or more. Tibial setae 5.3.3.3
EL WEE O CRO LEO banal Oa tate MORE bY Che MEW CURD Car aca Ch PCNAL Cac RCT COORCHICLS) ORCA B7CROTT TORONI- MEE RET 4
Trochanteral setae 1/0.0.0.0. Femoral setae 6.4.3.1. Tibial setae less than
Dra OtOnee tein reek see incase tarts sche chem RicreMan cade ete: ale cfae tate: etate, » (inv. ecpca 7
AMG) ow Eh CLOUOLALE BELAY OVA Gsald Mra khs i Atcha RMA et devia: tAMMENS c ceon Ns. o ctol ee pla ote aledely bho ote. ae 5
Wemoral saraceAis:3: wariceasrecht ins cckea diana toes o eyskaie tia di’ SUCGE Ra cOhole-a%> d:dtnias 6
5 (4). Coxal setae slender, parallel-sided, about twice as long as diameter of their
alveoli. Solenidion on tarsus II set in depression, but free. Posteroventral
margin of basis capituli uniformly sclerotized .......... zosteropis Fain
Coxal setae short, clavate, barely as long as diameter of their alveoli.
Solenidion on tarsus II concealed in narrow invagination. Sclerotization of
posteroventral margin of basis capituli interrupted medially ............
HEISE. Perdana Rd Meee OUTS anptarcha ctbetorstete ae. tee Bruns “Boyda)
Gn(4) allyes absent. [Coxalisetae 2s ai0) are. ccs seni siamie. sisieieiel: ted eis spatulata Fain
Ves mpresenitn mCOxalmSetaceaplica: Omm cm cr en-) sumer. /e.eeracbe cisrs myzomelae, D. sp.
7 (8). Tibial setae 5.3.2.3........... crassipes (Berlese and Trouessart), n. comb.
Tibiale setae s4sacatae vce sat sete eke ik EE Ree Rae ea. eee ate val maluri, n. sp.
BoypDAIA STURNI (Boyd)
Speleognathus sturni Boyd, 1948, Proc. ent. Soc. Wash., 50: 9; 1951,
J. Parasit., 37: 79 (where the author lists the locality of some Indian
specimens in the form of a binomen allegedly identifying one of the mynas) ;
nec Porter and Strandtmann, 1952, Tew. J. Sci., 4: 394. Boydaia sturm,
Fain, 1961, Bull. Annls Soc. r. ent. Belg., 97: 57; 1963, Bull. Inst. r. Set.
nat. Belg., 39: 56; Z. ParasitKde, 22: 371.
Previous records (both introduced Sturnidae, Passeriformes) .—Starling,
Sturnus vulgaris Linnaeus. Common myna, Acridotheres tristis (Linnaeus).
New Australian record.—S. vulgaris, Esk, 6.x.1966, R. D. and J. S. W.
(13 adults, 4 larvae).
BoOYDAIA ZOSTEROPIS Fain
Boydaia zosteropis Fain, 1963, Bull. Inst. r. Sci. nat. Belg., 39: 43.
This species, previously known only from an African silvereye, may now
be recorded from Australia: 2 adults and 2 larvae from a grey-backed silver-
eye, Zosterops lateralis (Latham) (Zosteropidae, Passeriformes), Mt. Nebo,
17.xi.1965, E. H. D.
BoyDAIA HIRUNDOAE Fain
Boydaia hirundoae Fain, 1956, Annls Parasit. hum. comp., 31: 661;
1958, Revue Zool. Bot. afr., 58: 180; 1968, Bull. Annls Soc. r. ent. Belg.,
99: 179; Domrow, 1965; Acarologia, 7: 43.
Previous records (both MHirundinidae, Passeriformes).—Welcome
swallow, Hirundo neoxena Gould, Brisbane. Common swallow (vagrant),
H. rustica Linnaeus.
New host record.—Fairy martin, Hylochelidon ariel (Gould) (Hirun-
dinidae), Windorah, 20.1.1966, R. D., D. J. M., and J. S. W. (1 larva).
BoypatlA FALCONIS Fain
Boydaia falconis Fain, 1956, Annls Parasit. hum. comp., 31: 657.
This African species is now known also to occur in Australia: 35 adults
and 1 larva from brown hawks, Falco berigora Vigors and Horsfield
(Falconidae, Falconiformes), Kowanyama, 27.x and 9.xi.1965, R. D.
BoyYDAIA SPATULATA Fain
(Figs. 223-226)
Boydaia sturni var. spatulata Fain, 1955, Annis Soc. belge Méd. trop.,
35: 695. See Art. 45. Boydaia spatulata, Fain, 1956, Riv. Parassit., 17: 33;
1956, Revue Zool. Bot. afr., 53: 43; 1956, Annls Parasit. hum. comp., 31: 651.
382 THE NASAL MITES OF QUEENSLAND BIRDS
Fain (1963c) has detailed a group of five African and European species
centred around B. spatulata, which, while distinguishable as larvae, are
scarcely separable as adults. In the absence of immatures, I now record
Fain’s species from three new Australian hosts (Meliphagidae, Passeri-
formes): 19 from a banded honeyeater, Myzomela pectoralis Gould, Kowan-
yama, 23.x.1965; 1 2 from a noisy friar-bird, Philemon corniculatus (Latham),
Esk, 5.x.1965; and 192 from a little friar-bird, P. citreogularis (Gould),
Charleville, 14.1.1966, R. D., D. J. M., and J. S.-W.
The specimen illustrated is from Myzomela Vigors and Horsfield, and
has .the following measurements: idiosoma 410u long, 3144 wide; capitulum
93 long, 82» wide; palpi 35y long; leg I 315, II 266, III 249, IV 270» long.
Setal formulae for legs: coxae 2.1.1.0; trochanters 1.1.0.0; femora 7.4.3.3;
genua 4.4.3.5; tibiae 5.3.3.3; tarsi 12.8.7.7. The specimens from Philemon
Vieillot are considerably larger, but show identical chaetotaxy, although,
in fact, the following minor variants occur: genital setae 5.4 (Myzomela) ;
hypostomal setae 2.1 (P. corniculatus); femora I-II 7.4/6.3, genu II 4.3,
and tibia IV 3.4 (P. citerogularis—some broken cuticle from the body is
Wy 224
a vA
| J
n
XR nN
A i
Lee
& %
Figs 223-225. Boydaia spatulata Fain.—223, Dorsal view of adult from Myzomela
pectoralis. 224-225, claws and pulvilli of adults from WM. pectoralis and _Philemon
citreogularis (at twice indicated scale).
ROBERT DOMROW 383
folded over one leg IV, but the four setae in question appear to lie beneath
it, and, therefore, to belong to the tibial, and not to the idiosomal series,
Some measurements are: capitulum 116 long, 103” wide; palpi 54» long;
leg II 350-865, ITT 320-840, IV 345-375» long (excluding ambulacra, but
including coxae).
The sensillae in the series from Myzomela are 32» long and about one-
tenth as broad; in specimens from Philemon, they are 48-45» long and
noticeably more slender (one-fifteenth as broad as long).
Fig. 226. Boydaia spatulata Fain—Ventral view of adult from Myzomela pectoralis.
BoyDAIA MYZOMELAE, N. Sp.
(Figs. 227-229)
Diagnosis.—B. myzomelae is related, on chaetotaxy, to the five species
centred around B. spatulata (see Fain, 1963c, and discussion on that species
above), but may easily be separated from them by the coxal formula (2.1.2.0,
not 2.1.1.0) and the presence of distinct eyes.
Two other species of Boydaia, B. hirundoae and B. psalidoprocnei Fain,
show a coxal formula 2.1.2.0, but are distinct from B. myzomelae both
morphologically (external seta on coxa I minute) and ecologically (hosts
swallows, Hirundinidae, Passeriformes) (see Fain, 1963c).
384 THH NASAL MITES OF QUEENSLAND BIRDS
Types.—Holotype female from a scarlet honeyeater, Myzomela sanguino-
lenta (Latham) (Meliphagidae, Passeriformes), Esk, 6.x.1966, R. D. and
J. S. W. Holotype N. I. C.
Female.—Idiosoma engorged, ruptured during mounting procedure, 465u
long, 3154 wide across humeral prominences. Cuticle finely striate-punctate.
WMH) WZ
\\
x
R w
We
: RD
Figs 227-228. Boydaia myzomelae, n. sp. (adult from Myzomela sanguinolenta) —227,
Dorsal view. 228, Pulvillus (at twice indicated scale).
Kyes sharply outlined, with corneae (Fig. 227). Sensiilae 32. long, very
slightly clavate. Presensillary setae small, 2 in number. Postsensillary setae
3.4.2.2.4.2.
Sternal setae 2+2.2. Genital setae 2.3/3.2 Anal setae 2.2 (Fig. 229).
Legs with reticulate subcuticular markings, I 302, II 275, III 250» long
(IV lacking except for coxae and one trochanter). Setal formulae: coxae
2.1.2.0; trochanters 1.1.0.0; femora 7.4.3; genua 4.4.3.-; tibiae 5.3.3.-; tarsi
ROBERT DOMROW BRD
12.8.7. Tarsi I and II with solenidion; tibia I with invaginated sensory
organ.” Claws equal, of uniform diameter except at tip, roughened internally
near basal angulation; retractable into dorsodistal pits in tarsi. Pulvilli
directed upwardly between claws, bifid, with each arm expanded distally and
flared externally; with many filaments on plantar surface (Fig. 228).
Fig. 229. Boydaia myzomelae, n, sp.—Ventral view of adult from Myzomela sanguinolenta.
Capitulum 80» long, 90% wide, reticulate; hypostomal setae 2.2. Palpi
3-segmented, 40u long; setal formula 0.0.3; solenidion present on tarsus.
All setae barbulate (many on dorsum of idiosoma and legs appear, at
lower magnification, to terminate in hyaline expansion, which, however,
under oil-immersion, is seen to comprise merely shorter terminal setules) ;
Six setae on tarsus I and one on tarsus II with rod-like, hyaline extension
apically.
* These organs are treated more fully by Fain (19620). They are reminiscent
of the chemoreceptors of insects classified as sensilla coeloconica by Imms (1957).
386 THE NASAL MITES OF QUEENSLAND BIRDS
BoyDAIA CRASSIPES (Berlese and Trouessart), n. comb.
Ereynetes crassipes Berlese and Trouessart, 1889, Bull. Biblioth. scient.
Ouest, 2: 141. Speleognathus sturm Porter and Strandtmann, 1952, Tex. J.
Sci., 4: 394, nec Boyd, 1948, Proc. ent. Soc. Wash., 50: 9. See Fain (19586).
Boydaia nigra Fain, 1955, Annls Soc. belge Méd. trop., 35: 695; 1956, Riv.
Parassit., 17: 82; Revue Zool. Bot. afr., 53: 44; Annls Parasit. hum. comp.,
31: 658; 1957, Revue Zool. Bot. afr., 55: 254; 1958, Ibid., 58: 179; 1962,
Bull. Annis Soc. r. ent. Belg., 98: 266; 1968, Z. ParasitKde, 22: 371; Bull.
Annls' Soc. r. ent. Belg., 99: 179. New synonymy.
Previous records (all Passeriformes).—Yellow wagtail (vagrant),
Motacilla flava Linnaeus (Motacillidae). Goldfinch (introduced), Carduelis
carduelis (Linnaeus) (Fringillidae). House-sparrow (introduced), Passer
domesticus (Linnaeus) (Fringillidae).
New host record.—White-winged triller, Lalage tricolor (Swainson)
(Campophagidae, Passeriformes), Charleville, 14.1.1966, R. D., D. J. M., and
J. S. W.; Kowanyama, iv.1965, R. D. (33 adults, 9 larvae).
I accept the synonymy suggested by Fain (1957c). As in the case of
Ptilonyssus niteschi above, I believe this procedure preferable to accepting
both a nomen dubium and a later taxon which is almost certainly its junior
synonym. It should be confirmed by the formal designation of a neotype
when the group is next revised (Art. 75). Neither Lombardini (1936) nor
Fain (1964e), in their listings of the ereynetids in the Berlese collection in
Florence, mention this species, and Dr. M. H. Naudo tells me there is none in
the Trouessart collection in Paris. As with all these mites, the true locality
is the nasal passage of their specific host wherever it may exist, rather than
any fixed geographical area, so I see no objection to choosing one of Porter
and Strandtmann’s specimens, providing they are in good condition.
BoyDAIA MALURI, 0. sp.
(Figs 230-2385)
Diagnosis—B maluri falls among those small species of Boydaia with
a considerably reduced leg setation (Fain, 1963c), and may best be compared
with B. trochila Fain (1958a). However, the two species may be separated
by the setation (3, not 2 or 1) of genu IV, and the presence or absence of
a solenidion on the palpal tarsus. Lesser differences appear to lie in the
claws and pulvilli.
The host relationships should also be noted. B. maluri is from peculiarly
Australian “wrens” (Sylviidae, Passeriformes), while B. trochila was
described from hummingbirds, which belong to the neotropical family
Trochilidae (Apodiformes).
T'ypes—Holotype female from a lovely wren, Malurus amabilis Gould
(Sylviidae, Passeriformes), Cowley Beach, 9.viii.1965, R. D. and J. S. W.
Three females one of which is designated paratype) from a red-backed wren,
M. melanocephalus (Latham), Kowanyama, 22.xi.1965, R. D. Holotype
N. I. C.; paratype R. D.
Female.—Idiosoma (Fig. 231) engorged and pigmented in all specimens,
with humeral prominences; from 874x308, through 432x345 (figured), to
528x885 depending on degree of compression during mounting procedure.
Cuticle minutely striate-punctate. Eyes absent. Sensillae 24-27» long, very
slightly clavate. Presensillary setae minute, 2 in number. Postsensillary
setae 4.4.2.2.4.2.
Sternal setae 2+2.2. Genital setae 2.3/3.2 Anal setae 2.2 (Fig. 230).
ROBERT DOMROW 387
Legs (Figs. 284-285) with reticulate subcuticlar markings, 213, 193,
186, and 205» long in two specimens figured. Setal formulae: coxae 2.1.1.0;
trochanters 1.0.—— (M. amabilis), 0.0.0.0 (M. melanocephalus) ; femora 6.4.3.1;
genua 4.4.3.8; tibiae 4.2.2.2; tarsi 12.8.7.7. Tarsi I and II with solenidion;
—
q
v
s
°
NX
DNAS
6
R
VS RD
Figs 230-232. Boydaia maluri, n. sp. (adult from Malurus melanocephalus) —230-231,
Ventral and dorsal views. 232, Pulvillus (at twice indicated scale).
Figs 233-235. Boydaia maluri, n. sp. (adult from Malurus amabilis) —233, Ventral view
of capitulum. 234-235, Ventral and dorsal views of leg I.
388 THH NASAL MITES OF QUEENSLAND BIRDS
tibia I with invaginated sensory organ. Claws equal, simple, of uniform
diameter except at tip; retractable into dorsodistal excavations in tarsi.
Pulvilli (Fig. 282) directed upwardly between claws, apparently entire
except for weak distal cleft; with numerous filaments on plantar surface.
Capitulum (Fig. 283) 80 long, 854 wide, reticulate; hypostomal setae
2.2. Palpi 3-segmented, 36 long; setal formula 0.0.3; solenidion present on
tarsus.
All setae barbulate; six on tarsus I and one on tarsus II with expanded,
hyaline extension distally.
Genus Nerosoypara Fain
Neoboydaia Fain, 1958, Revue Zool. Bot. afr., 58: 177. Type-species
Boydaia philomachi Fain, 1956, Riv. Parassit., 17:27. Ralliboydaia Fain,
1962, Revue Zool. Bot. afr., 66: 365. 'Type-species Neoboydma (Ralliiboydaia)
latiralli Fain, 1962, Loc. cit., 365. The original spelling is a lapsus calami
based on the host-genus Laterallus Bonaparte, and was subsequently corrected
to lateralli under Art. 32 by Fain (19646). Aureliania Fain, 1958, Revue Zool.
Bot. afr., 58: 177. Type-species Boydaia aureliani Fain, 1955, Annls Soc. belge
Méd. trop., 35: 694.
Key to adults of Australian species of NEOBOYDAIA
1. Presensillary setae minute. Palpi with two segments .................. 2
Presensillary setae normal. Palpi with ome segment .................. 3
2 (1). Genital setae 6.6. Anal setae 1.1. Coxal setae 2.1.1.0. Femoral setae 5.4.3.2.
alpailisSebacwiO Ser tok sihees bc. caiduoe suse Wevew sc ces ccush one ence aE eee philomachi Fain
Genital setae 4.4. Anal setae 2.2. Coxal setae 2.1.1.1. Femoral setae 4.3.2.2.
Walipal’ Setaer Wig OFF. ee Ge he ans ee See a ene oN psittaculae Fain
3 (1). First pair of postsensillary setae present. Sensillae filamentous. Hypostomal
UCI fy Jon, See ne ee ae Pee REET nab oo o.0000000 4
First pair of postsensillary setae absent. Sensillae globose. Hypostomal
SOtae HOlO nee Se icdey sah eucio sche Civile Din erat ae Tee eee merops (Fain)
4 (3). Sternal setae 2.2.0. Coxal setae 1.1.1.1. Trochanteral setae 0.0.0.0. ..........
AES CIRCE OEY Cite Oa ERR oe END eT CEG Oe eect OCS o aureliani (Fain)
Sternal setae 2.2.2. Coxal setae 2.1.1.1. Trochanteral setae 1.1.0.0............
Be OR EGE FO. ORO Cac RCH AICRC TE COCCI COE MOREE CR nor hac bacarion ic ato colymbiformi Clark
NEOBOYDAIA PHILOMACHI (Fain)
Boydaia philomachi Fain, 1958, Riv. Parassit., 17: 27; 1964, Annls Mus.
r. Afr. cent. Sér. 8vo, 132: 141; Clark, 1964, J. Parasit., 50: 160; Domrow,
1965, Acarologia, 7: 48.
Previous records (all Scolopacidae, Charadriiformes) .—Bar-tailed godwit,
Inmosa lapponica (Linnaeus), Half Tide. Wood-sandpiper, Tringa glareola
Linnaeus. American pectoral sandpiper, Hrolia melanotos (Vieillot). Ruff
(vagrant), Philomachus pugnagz (Linnaeus).
NEOBOYDAIA PSITTACULAE Fain
Neoboydaia psittaculae Fain, 1962, Bull. Annls Soc. r. ent. Belg., 98:
318.
Previous record.—Peach-faced lovebird (introduced cage-bird), Agapornis
roseicollis (Vieillot) (Psittacidae, Psittaciformes).
New host records (both Psittacidae).—Rainbow-lorikeet, Trichoglossus
moluccanus (Gmelin), Kowanyama, 1.xi.1965, R. D. (1 adult). Varied
lorikeet, Psitteuteles versicolor (Lear), Kowanyama, 3.xi.1965, R. D. (1 adult).
ROBERT DOMROW 389
NEOBOYDAIA AURBLIANI (Fain)
Boydaia aureliani Fain, 1955, Annls Soc. belge Méd. trop., 35: 694; 1956,
Revue Zool. Bot. afr., 53: 36. Boydaia (Aureliania) aureliani, Fain, 1958,
Revue Zool. Bot. afr., 58: 177. Neoboydaia (Aureliania) aureliani, Fain,
1963, Bull. Annls Soc. r, ent. Belg., 99:181; Bull. Inst. r. Sci. nat. Belg.,
39: 52.
Previous record.—Barn-owl, Tyto alba (Scopoli) (Tytonidae, Strigi-
formes). ’
NzOBOYDAIA MmROPS (Fain)
13 J =4
Boydaia merops Fain, 1955, Annls Soc. belge Méd. trop., 35: 694; 1956,
Riv. Parassit., 17: 30; Domrow, 1966, Proc. Linn. Soc. N.S.W., 90: 210.
Previous record.—Rainbow-bird, Merops ornatus Latham (Meropidae,
Coraciiformes), Esk and Innisfail. Also Kowanyama.
NEHOBOYDAIA COLYMBIFORMI Clark
Neoboydaia colymbiformi Clark, 1964, J. Parasit., 50: 158.
This species may now be recorded from Australia: 1 adult from a little
grebe, Podiceps ruficollis (Vroeg) (Podicipidae, Podicipiformes), Esk, 15.v.
1965, I. D. F. and J. S. W. The only previous record is from an American
grebe (Colymbus Linneaus, a name proscribed in favour of Podiceps Latham,
see Thomson, 1964).
Genus OPHTHALMOGNATHUS Dubinin
Ophthalmognathus Dubinin, 1957, Trudy leningr. Obshch. Estest., 73: 66.
Type-species Ophthalmognathus dogieli Dubinin, 1957, Loc. cit., 68. (—=Spele-
ognathus schoutedeni Fain, v. infra). Neospeleognathus Fain, 1958, Revue
Zool. Bot. afr., 58: 178. Type-species Speleognathus schoutedeni Fain, 1955,
Annls Soc. belge Méd. trop., 35: 695. New synonymy. Trispeleognathus
Fain, 1958, Revue Zool. Bot. afr., 58: 178. Type-species Speleognathus
striatus Crossley, 1952, J. Parasit., 38: 385. New synonymy. Metaboydaia
Fain, 1962, Bull. Inst. r. Sci. nat. Belg., 38: 7. Type-species Speleognathus
poffer Fain, 1955, Annls Soc. belge Méd. trop., 35: 695; 1956, Revue Zool.
Bot. afr., 53: 23. New synonymy.
The decision to accept Ophthalmognathus (see Fain, 19580) leaves one
only to note that, in the question of considering Neospeleognathus and
Trispeleognathus (both originally raised as subgenera of Speleognathus by
Fain, 1958b) as subgenera of a genus in its own right, Fain (1963c), as
first reviser, was free under Rec. 24 to choose the latter for the generic name,
although the former has precedence of position. Presumably, he was influenced
by his intermediate classification (1962d), in which Trispeleognathus was
given full generic rank, while Neospeleognathus was considered a subgenus of
Neoboydaia Fain.
Key to adults of Australian species of OPHTHALMOGNATHUS
i, Coxal setae 2.1.1.1. Femoral setae more than 5.4.3.2. Hypostomal setae 1.1.
Palpimwithechneesseamentsunseta tone Olea aia crete ote siertetelo) =) = = 2
Coxal setae 2.1.1.0. Femoral setae 5.4.3.2. Hypostomal setae 2.2. Palpi with
two segments, setation 0/0:2. ...-.......... schoutedeni Fain, n. comb.
2 (1). Genital setae 5.5. Femoral setae 5.4.3.3. ........ striatus (Crossley), n. comb.
Genital setae 4.4 Femoral setae 6.4.3.4 .................. accipitris, TD. sp.
OPHTHALMOGNATHUS STRIATUS (Crossley), n. comb.
Speleognathus striatus Crossley, 1952, J. Parasit., 38: 385; Fain, 1956,
Revue Zool. Bot. afr., 53: 18. Speleognathus (Trispeleognathus) striatus,
Fain, 1958, Revue Zool. Bot. afr., 58: 178. Trispeleognathus (Trispeleog-
nathus) striatus, Fain, 1963, Bull. Inst. r. Sci. nat. Belg., 39: 53. Trispele-
ognatus striatus (sic) do Amaral, 1963, Archos Inst. biol., S Paulo, 30:91.
390 THE NASAL MITES OF QUEENSLAND BIRDS
Previous record.—Domestic pigeon (introduced), Columba liwia Gmelin
(Columbidae, Columbiformes).
OPHTHALMOGNATHUS ACCIPITRIS, 1. Sp.
(Figs 236-237)
Diagnosis ——Three species of Ophthalmognathus s. s.* (te. with 3-seg-
mented palpi) are known. O. striatus and O. womersleyi (Fain), n. comb.,
are both listed from Africa and the Americas, from various pigeons (Columbi-
Fig. 236. Ophthalmognathus accipitris, n. sp.—Dorsal view of adult from Accipiter
fasciatus.
*It_ is a pity to have to note that acarological literature is becoming increasingly
crowded with misspellings of numerous terms in common taxonomic use. This one
and its partner are especially prone to suffer (sensu stricta, sensu latu, sensu latus),
but other examples are common, e.g. genus incertus, nomine nuda, species novum,
sp. novo, and status novum. There are also lapses in taxonomic names (dividus,
femuralis, flumenicola, inexspectatus, and omniphagous); in morphological terms
(endapodal, metatarsii, scutae, tarsae, and ventrum); and in general (a corrigenda,
atennuated, detritis, hiati, and in statuo quo). This unconcern for a language which
is both exact and the backbone of scientific terminology is inexcusable, and, regrettably,
even the Code (Stoll et al., 1964), in reducing Art. 31 to a mere Recommendation,
has taken a step towards the stand that “scientific names ... are meaningless since
Latin is no longer an effective scientific language’.
ROBERT DOMROW 391
formes) and ducks (Anseriformes), respectively (Clark, 1958; Fain, 1963c;
do Amaral, 19680). Full morphological details are given by Fain (1963c),
and O. accipitris, from an Australian goshawk (Falconiformes), may be
separated from both by the setal formulae of the genitalia (4.4, not 5.5) and
femora (6.4.3.4, not 5.4.8.3). O. accipitris further differs from O. womersleyi
in showing two pairs (not one) of anal setae, and lacking the single seta on
the palpal tibia. .
The third known species, from a Brazilian pigeon, is O. ewricoi (do
Amaral, 1965), n. comb. It is not as well characterized, but differs from all
: Ns
Fig. 237. Ophthalmognathus accipitris, n. sp—Ventral view of adult from Accipiter
fasciatus.
three species discussed above in showing a coxal formula 2.0.1.0 rather than
2.1.1.1, and two pairs of gnathosomal setae rather than one.
Types.—Holotype female and one paratype female from an Australian
goshawk, Accipiter fasciatus (Vigors and Horsfield) (Accipitridae, Falconi-
formes), Kowanyama, 25.x.1966, R. D. Holotype N. I. C.; paratype R. D.
Female.—Idiosoma 594 and 616 long in somewhat compressed condition.
Cuticle finely striate on body, minutely, but densely aciculate on appendages.
392 THE NASAL MITES OF QUEENSLAND BIRDS
Eyes present, but small, evident more by cessation of cuticular striae than
by any definite corneae. Scutum divided into two longitudinal fragments
(Fig. 236). Sensillae filamentous, simple. Dorsal setal formula (including
presensillary pair) 2.2(1).4.2.2.4.2, all barbulate (except exterior pair of
subposterior row, which are drawn out distally into filaments).
Sternal setae 2.2.1 in both specimens, but presumably 2.2.2 normally.
Genital setae 1.3/3.1; anal setae 2.2(1). All barbulate (Fig. 237).
Setal formulae for legs as follows: coxae 2.1.1.1; trochanters 1.1.0.0;
femora 6.4.3.4; genua 4.4.3.3; tibiae 5.3.3.3; tarsi 12.8.7.7. All barbulate
except as follows: one seta on anterior margin of femur I distinctly, and
occasional seta on some genua and tibiae slightly filamentous distally; tibiae
with 8.1.1.1 dorsal setae entirely filamentous; some rounded, barbulate setae
at tips of tarsi I-II with short, hyaline, apically expanded extensions. Tarsi
I-III each with dorsal solenidion. Tibia I with invaginated sensory organ.
Claws paired, subequal, retractable into dorso-distal excavations in tarsi.
Pulvilli entire, bearing filaments.
Capitulum and legs with distinct reticulate pattern beneath cuticle.
Palpi 3-segmented. Tibia with one seta dorsally; tarsus with one seta
dorsally, and two setae and solenidion ventrally. Only two fully formed
hypostomal setae present. All capitular setae barbulate, omitting vestiges
of second hypostomal pair present between normal pair. Chelicerae drawn
out distally into ventrally-directed stylets.
OPHTHALMOGNATHUS SCHOUTEDENI (Fain), n. comb.
Speleognathus schoutedeni Fain, 1955, Annls Soc. belge Méd. trop.,
35: 695; 1956, Revue Zool. Bot. afr., 53: 18. Ophthalmognathus dogieh
Dubinin, 1957, Trudy leningr. Obshch. Estest., 73:68. New synonymy.
A third species of this genus may also be recorded from Australia: 6
adults from nankeen night herons, Nycticoraz caledonicus (Gmelin)
(Ardeidae, Ciconiiformes), Esk and Kowanyama, 5.i.1966 and 6.iv.1965, R. D.
and J. S. W. The only previous records are African, NV. nycticorax (Linnaeus)
and Ardeola idae (Hartlaub).
I accept the synonymy suggested by Fain (19588). Both taxa were
described from herons.
-Genus SPELEOGNATHOPSIS Cooreman
Speleognathopsis Cooreman, 1954, Annls Parasit. hum. comp., 29 : 428.
Type-species Speleognathopsis galli Cooreman, 1954, Loc. cit., 429.
Key to adults of Australian species of SPELEOGNATHOPSIS
iL, Presensillary setae present. Four or more setae in first postsensillary row.
Anal setae 3.3. Coxal setal formula commencing 2.1.1 (seta on coxa II
of S. benoiti Fain minute). Femoral setae more than 5.4.3.2 ........ 2)
Presensillary setae absent. Two setae in first postsensillary row. Anal
setae 1.1 Coxal setae 1.0.1.0 Femoral setae 5.4.3.2 ...... galli Cooreman
2. (1). Scutum transverse. Seta on coxa II of normal size. Pulvilli simple. Palpal
SOCAC OLS S 1.t. teacaerey. atanayed aeokSPouie. chews setoa ea one hcg wmreiey Same tema porphyrionis Domrow
Scutum elongate. Seta on coxa II minute. Pulvilli bifid. Palpal setae 1.3
ERA Nelis yale Pebobave Sunes meh oye! Selsaulehsavlesieues shan Some penal s Vanes ewer te welran apa a eee eee benoiti Fain
SPELEOGNATHOPSIS GALLI Cooreman
Speleognathopsis galli Cooreman, 1954, Annls Parasit. hum. comp.,
29: 429; Fain, 1963, Bull. Inst. r. Sci. nat. Belg., 39: 53.
Previous records (both Galliformes).—Domestic fowl (introduced),
Gallus gallus (Linnaeus) (Phasianidae). Guinea fowl (introduced), Numida
meleagris (Linnaeus) (Numididae).
ROBERT DOMROW 393
Since writing this, I have seen 7 adults of this species, previously known
only from Rwanda, from the nasal sinuses and nares of domestic fowls in
villages in the Western Sepik District (near Irian Barat border), v.1966,
N. Talbot.
SPHrLEOGNATHOPSIS PORPHYRIONIS Domrow
Speleognathopsis porphyrionis Domrow, 1965, Acarologia, 7: 44.
Previous record.—Eastern swamphen, Porphyrio melanotus Temminck
(Rallidae, Gruiformes), Samford.
SPELEOGNATHOPSIS BENOITI Fain
Speleognathopsis benoiti Fain, 1955, Annis Soc. belge Méd. trop., 35: 696;
1956, Revue Zool. Bot. afr., 53: 29; Domrow, 1965, Acarologia, 7: 44; 1966,
Proc. Linn. Soc. N.S.W., 90: 210; McDaniel, 1967, Tex. J. Sci., 19: 94.
Speleognathopsis charadricola Fain, 1964, Revue Zool. Bot. afr., 70: 35.
Speleognathopsis wai Fain, Vercammen-Grandjean, and Wagner, 1966, Acta
zool. path. antverp., 41: 115. New synonymy.
I have already noted (19660) considerable intraspecific variation in this
parasite of charadriiforms, and commend to the reader Marshall’s statement
on taxonomy (in Parker and Haswell, 1962).
Previous records (all Charadriidae, Charadriiformes).— Red-kneed
dotterel, Hrythrogonys cinctus Gould, Kowanyama. Masked plover, Lobibyx
miles (Boddaert), Kowanyama. Grey plover, Squatarola squatarola
(Linnaeus). Black-fronted dotterel, Charadrius melanops Vieillot, Esk. Also
Kowanyama.
Family EPIDERMOPTIDAE
Subfamily TURBINOPTINAE
This latter group has usually been accorded full familial status in recent
years, but this period has also seen the erection of annectant taxa such as
the Heterocoptidae (Fain, 1967a). I therefore return the group to its
original family, the Epidermoptidae, noting that, while the epidermoptines
are skin-parasites of birds, the turbinoptines inhabit the outermost parts
of the nasal cavity. Such a treatment is parallel to that accorded the other
two groups of nasal parasites of birds discussed above.
Even though I have used the characters of the female genitalia and the
male clasping apparatus in the primary dichotomies in the following key to
genera, it should be noted that the groups so obtained are not correlated.
For example, while females of Rhinoptes de Castro and Pereira and
Schoutedenocoptes Fain both key out through the second half of couplet 2,
males of the former diverge from those of the latter at couplet 7. Nor do
the two genera show similar tarsal modifications, yet both parasitize
phasianoids (Galliformes).
Key to Australian genera of TURBINOPTINAE
1. NOI AT OS Ms see ecie eave Here ee a eee ec ms elite Sie abene Se sre cies -e ece suele 2 Se eis 2
IY IETS AS ECS Aah OE Ot be oles Cr Dic CL GLCROTE OIE) ES CRESCENT CLEC OES CRCRCICNET ETT eon Io oar 7
2G) ceendosyniumsabsentsmaViUlvamtGansVverse) cel -scices aecleseciie sacle ce 1 sie 3
Endogynium present. Vulva in form of inverted Y .....................--- 4
Se 02) eral starsic within twiowiclawSt vs ant css ote oc een clear cs nclee Turbinoptes Boyd
All@tarsi*withivone iclawiase. 1o42 acme oo eek bre bade Passerrhinoptes Fain
401 (2) Pee Als tarsiawithetwo (claws 4 sis «acs tec tin Sie ws clersl als «6 aes sete Ozleya Domrow
All tarsi pnotawath two Claws! Men acs sets cis-elshsle ie stave elvis gue Nel els srelclere eteielers iors ore 5
5 (4). All tarsi very short and heavily sclerotized, with at least one claw ...... 6
Only tarsi-I-II very short and heavily sclerotized. Tarsi I-II with one claw.
Tarsi- (-nve without claws 220---s52escc0+--0.6-- Schoutedenocoptes Fain
394 THE NASAL MITES OF QUEENSLAND BIRDS
6 (5). Tarsi I-IL with two claws. Tarsi III-IV with one clawtihve laos aed eee
al ia Pegs, Pte ged AER Oe i hea loan Mi Un Caracas Rhinoptes de Castro and Pereira
AIDE tarsiswith One (Clawalis . mice cine nts). plete nt clerks avschatltel stabil Mycteroptes Fain
7 (1). Body rounded posteriorly. Anal suckers minute ................0.ese00. 8
Body bilobed posteriorly. Anal suckers distinct .................0.secee 10
8 (7). Tarsi I-II with two claws. Tarsi IIIJ-IV with one claw ....................
AT] tarsi: with ome: Claws bs. cic scutud So cnchoutes nla Sh cus Gr Ate oes ae hes dee ee oe )
OCS) “Mpinieral W “Eee aici te erate etecee (oped c ccehekens 4 Rott SRSA coal HONS Mycteroptes Fain
Mpimerawlyfused 4c tea. a i ARIAS: IO Ae 2S ae Passerrhinoptes Fain
10 (7). All tarsi with two claws. Legs III-IV Send s) sis) vie) sha (abe skelep-f-1 espana 11
Tarsi I-II with one claw. Tarsi III-IV without claws. Leg IV considerably
smallervthaneil dl: Ais are eee cee ets Bis, Ae bone unseat Schoutedenocoptes Fain
11 (10). Dorsum unarmed except for minute propodosomal shield .... Oxleya Domrow
Dorsum with extensive hysterosomal shield in addition to propodosomal
SHUCLG Ss cieate apse tvanieis tee thaceke ene cd oLekeaee i awe oe eee Turbinoptes Boyd
Genus TuRBINOPTES Boyd
Turbinoptes Boyd, 1949, J. Parasit., 35: 295. Type-species Turbinoptes
strandtmanni Boyd, 1949, Loc. cit., 295.
TURBINOPTES STRANDTMANNI Boyd
Turbinoptes strandtmann Boyd, 1949, J. Parasit., 35: 295; Fain, 1956,
Revue Zool. Bot. afr., 54: 222; 1960, Ibid., 62: 101; 1962, Bull. Annls Soc.
r. ent. Belg., 98: 266; 1963, Ibid., 99: 179; Dubinin and Snegireva, 1957,
Zool. Zh., 36: 204; Domrow, 1965, Acarologia, 7: 90.
Previous records (all Charadriiformes).—Silver gull, Larus novaehol-
landiae Stephens (Laridae), Half Tide. Whimbrel, Numenius phaeopus
(Linnaeus), Scolopacidae. Common sandpiper, Tringa hypoleuca Linnaeus
(Scolopacidae).
New host record——Eastern curlew, Numenius madagascarensis (Lin-
naeus), Tin Can Bay, 22.viii.1966, R. D. and J. S. W. (829 9, 174 4, 6
nymphs).
New Australian record.—N. phaeopus, Tin Can Bay, 26.x.1966, R. D.
and J. S. W. (72 2,52 ¢).
Genus PASSERRHINOPTES Fain
Passerrhinoptes Fain, 1956, Revue Zool. Bot. afr., 54: 216. Type-species
Passerrhinoptes andropadi Fain, 1956, Loc. cit., 217; 1957, Annls Mus. r.
Congo belge Sér. 8vo, 60: 30.
PASSERRHINOPTES POMATOSTOMI Domrow
Passerrhinoptes pomatostomi Domrow, 1966, Proc. Linn. Soc. N.S.W..,
90: 213.
Previous record.—Grey-crowned SNES, Pomatostomus temporalis
(Vigors and Horsfield) (Timaliidae, Passeriformes), Esk. Also Condamine.
Genus OxLEYA Domrow
Ozleya Domrow, 1965, Acarologia, 7:85. Type-species Oxleya podargi
Domrow, 1965, Loc. cit., 85.
OxLHyA PopARGI Domrow
Oaleya podargi Domrow, 1965, Acarologia, 7: 85.
Previous record.—Tawny frogmouth, Podargus strigoides (Latham)
(Podargidae, Caprimulgiformes), Samford and Cobble Ck. Also Hsk.
ROBPRT DOMROW 395
Genus ScuoutTepenocorres ain
Schoutedenocoptes Fain, 1956, Revue Zool. Bot. afr., 54: 210. Type-species
Schoutedenocoptes numidae Fain, 1956, Loc. cit., 211.
ScHOUTEDENOCOPTES NUMIDAR Fain
Schoutedenocoptes numidae Fain, 1956, Revue Zool. Bot. afr., 54: 211;
1957, Annls Mus. r. Congo belge Sér. 8vo, 60: 17.
Previous record.—Guinea fowl (introduced), Numida_ meleagris
(Linnaeus) (Numididae, Galliformes).
Genus Rutnopres de Castro and Pereira
Rhinoptes de Castro and Pereira, 1951, Archos Inst. biol., S Paulo, 20:
67. Type-species Rhinoptes gallinae de Castro and Pereira, 1951, Loc. cit.,
67.
RHINOPTES GALLINAB de Castro and Pereira
Rhinoptes gallinae de. Castro and Pereira, 1951, Archos Inst. biol., 8
Paulo, 20: 67; Domrow, 1965, Acarologia, 7: 90. Rhinoptes pternistis Fain,
1956, Revue Zool. Bot. afr., 54: 221; 1957, Annls Mus. r. Congo belge Sér.
8vo, 60: 37. New synonymy.
Previous records (both Phasianidae, Galliformes) Domestic fowl
(introduced), Gallus gallus (Linnaeus). Brown quail, Synoicus australis
(Latham), Dayboro and Chelona.
I would confirm here the synonymy implicit in an earlier note (Domrow,
19656).
Genus Mycrteroptses Fain
Mycteroptes Fain, 1956, Revue Zool. Bot. afr., 54: 219. Type-species
Mycteroptes basilewskyi Fain, 1956, Loc. cit., 220.
MYCTEROPTES BASILEWSKYI Fain
Mycteroptes basilewskyi Fain, 1956, Revue Zool. Bot. afr., 54: 220; 1957,
Annls Mus. r. Congo belge Sér. 8vo, 60: 33.
This genus and species, previously known only from an African roller
of the genus Coracias Linnaeus may now be recorded from Australia:
17 2 2,14, and 6 nymphs from an eastern broad-billed roller, Hurystomus
orientalis (Linnaeus) (Coraciidae, Coraciiformes), Esk, 5.x.1965, R. D. and
J. 8S. W.
V. HOS?T-SPECIFICITY AND ZOOGEOGRAPHY
At the beginning of this study, I was surprised to find so many species.
originally described from overseas birds, also present in Australia, but it
soon became clear that this was due simply to the fact that these mites
had never been searched for in Australia. Thus only 39 (39%) of the 99
rhinonyssines, four (21%) of the 19 speleognathines, and two (33%) of the
six turbinoptines now known (or expected to occur) in Australia have proved
to be new species. This represents an overall figure of 36% (45 out of 124
species). In the following discussion on the host-specificity and zoogeography
of the Australian fauna, the species are discussed, as far as possible, in
systematic order. Those I have described as new since the beginning of this
study are indicated by an asterisk, as it is interesting to consider to what
extent the hosts of these species are restricted to the Australian zoogeo-
graphical region.
396 THE NASAL MITES OF QUEENSLAND BIRDS
(a) Tinaminyssus
The three species from psittacids are the only Australian members of
this genus to retain the tritosternum. This structure is, however, in the
process of regression, and is absent in other species from extra-Australian
psittacids, e.g. T. baforti (Fain), n. comb., and 7. phalliger (Fain), n. comb.
(see Fain, 1963e, 19650); Domrow, 1964a; Wilson, 1964, 19666). The
opisthonotal shield is also regressing in this group. The host-specificity of
these three species is discussed in detail above in Section IV under
T. aprosmicti. In essence, each has several hosts within well defined psittacid
groups: JT. kakatuae* parasitizes crested species (cockatoos and the
cockatiel), J’. aprosmicti* various parrots, and T'. trichoglossi* various
lorikeets. However, although the Psittacidae (as well as the Columbidae
and Alcedinidae, v. infra) have undergone considerable radiation in the
Australian region, all three families are virtually cosmopolitan, and some of
their newly described parasites have already been recorded in New Guinea,
Indonesia, and the Philippines (Wilson, 1964, 1966a, b, 1968a, 6). There is
no reason therefore, not to expect them even further afield.
T. belopolskii has been recorded from all the continents except South
America, and is restricted to the Ardeidae.
T. epileus is, as yet, known only from New Guinea, but it parasitizes
a falconid which extends well into Australia.
The largest series (11 species) of Australian species of Tinaminyssus
is that restricted to columbids. The simpler species, 7.e. those lacking
basally inflated setae on the legs and venter, may be divided into two groups
depending on the postanal seta, yielding the sequences—(7) postanal seta
present: 7’. melloi, T. ocyphabus* and T. hirtus; and (ii) postanal seta absent:
T. columbae, T. megaloprepiae,* T. geopeliae, T. ptilinopi, and T. myristici-
worae.* The widespread J. mellot occurs in all three introduced ground-
pigeons (Columbinae), which are primarily seed-eaters. Its remaining two
hosts (Columba norfolciensis and Leucosarcia melanoleuca) are jungle-dwell-
ing, seed- and fruit-eating members of the same subfamily. 7’. ocyphabus is
known only from the peculiarly Australian Ocyphaps Gray, while T. hirtus
is known from Chalcophaps Gould in the Philippines, New Guinea, and Aus-
tralia. These are again columbines, the former typical of open forest, the latter
of rain-forest. T. columbae is also widespread in the domestic pigeon, and has
been once recorded from Ocyphaps, while T. geopeliae is restricted to Geopelia
Swainson in Malaysia, the Philippines, New Guinea, Australia, and Hawaii
(introduced). The remaining three species of the sequence are parasites
of jungle-dwelling fruit-pigeons (Treroninae). 7. ptilinopi has been recorded
from Sphenurus Swainson in Taiwan, and is widespread from S. E. Asia to
New Guinea in Ptilinopus Swainson, one of the known hosts extending well
into Australia. 7. megaloprepiae and T. myristicivorae are known only from
their original Australian hosts.
The second group comprises JT. phabus,* recorded only from Phaps Selby,
an Australian columbine of the open forest, 7. macropygiae, restricted to
a jungle-dwelling columbine (Macropygia Swainson) in the Philippines,
New Guinea, and Australia, and 7. welchi,* a parasite of a treronine. Clearly,
there is a tendency for the more advanced species of this genus to be found
parasitizing the jungle-dwelling treronines rather than the more typical
columbines.
There remain two species from alcedinids: 7. haleyonus* is common
in Halcyon Swainson in the Philippines, New Guinea, and Australia, while
T’. daceloae* has only been recorded from Dacelo, an Australian genus.
ROBERT DOMROW 397
(b) Larinyssus
Of the two known species, L. orbicularis is cosmopolitan in the Laridae,
and L. benoiti has been recorded from African and Australian glareolids.
(c) Rallinyssus
This compact genus is cosmopolitan and restricted to rallids. Of the
two species without circumanal frills, Rk. amaurornis is known from two
hosts in Taiwan and the Philippines, and R. congolensis from a further
two in Rwanda and the Philippines. The range of one host of each species
extends into Australia. Of the two species with circumanal] frills, R. gallin-
ulae has been recorded from Belgium, New Guinea, and Australia, and
R. caudistigmus from the U.S.A., South Africa, New Guinea, and Australia.
Host-specificity is again low, the former species having been recorded from
five genera, and the latter from three of the same five (Wilson, 1965, 1967).
(d) Rhinonyssus
This genus is confined to water birds of several types, and the primary
dichotomy in the key in Section IV above is based on the work of Strandtmann
(1959). Of the species with a definite anal shield and compacted palpi,
R. rhinolethrum is cosmopolitan in a very wide range of anseriform genera,
while R. poliocephali is known from a single species of Podiceps Latham
in both Africa and Australia. Of the species with the anal shield obsolescent
and normal palpi, both R. coniventris and R. minutus are cosmopolitan
parasites of a wide range of charadriiforms, while R. sphenisci has only been
recorded from Sphenisciformes.
(e) Ruandanyssus
Of the two known species, R. terpsiphonei has a wide range of African
and Australian passeriform hosts, while R. artami* is as yet known only
from Australian artamids. However, the latter may be expected to coexist
with its host-family, which extends to S. E. Asia.
(f) Rhinoecius
This genus comprises several species, all restricted to the Strigiformes.
Of the Australian species, R. cooremani, originally recorded from Strix
Linnaeus in the U.S.A., here parasitizes Ninox Hodgson (also a strigid), while
R. tytonis probably coexists with the widespread barn-owl, Tyto alba
(Tytonidae).
(g) Ptilonyssus
Apart from three species noted below (P. triscutatus, P. nitzschi, and
P. cerchneis), the 53 Australian members of this genus are restricted to the
Passeriformes, and may conveniently be split into two major divisions
depending on whether or not the opisthonotal shield is reduced merely to
a pygidial complex. (The presence of metasternal setae is often, but not
always correlated with this division.) The first division may again be once
divided. Firstly, a group may be commenced with P. maluri* and P. colluri-
cinclae* (parasites of peculiarly Australian sylviids and pachycephalids, two
taxa often considered only as muscicapid subfamilies, see Thomson, 1964),
which lead by way of P. capitatus (a widespread parasite of alaudids) to
P. carduelis, P. emberizae, P. pygmaeus, P. hirsti, and P. neochmiae* (all
parasites of fringillids and ploceids, two widespread families of small grani-
_ vores). (P. neochmiae lacks peritremes, but this minor morphological point
398 THE NASAL MITES OF QUEENSLAND BIRDS
should not be allowed to override an otherwise convincing unity.) The group
closes with P. struthideae* and P. corcoracis,* two species lacking a cribrum
and parasitizing two monotypic, and peculiarly Australian corvid genera.
They appear closely related to P. nucifragae, a parasite of Nucifraga Brisson,
a palaearctic member of the same family.
The second group, involving increasing fragmentation and reduction of
the opisthonotal shield, includes seven species. In P. triscutatus, P. diop-
trornis, P. dicruri, and P. sittae, the pygidial shield is discrete. The first
is a parasite of meropids (Coraciiformes), while the other three parasitize
muscicapids (including turdids), dicrurids, and sittids (including certhiids),
respectively. All are widespread in Africa, Eurasia, and Australia.
P. ailuroedi,* an aberrant species with considerable mesonotal modifications,
probably also belongs here. Its hosts belong to the Ptilonorhynchidae, a
peculiarly Australian family near the end of the passerine series. On the
other hand, P. elbeli and P. angrensis both lack a pygidial shield, the former
being a parasite of Eurasian sturnids, while the latter parasitizes the cosmo-
politan family Hirundinidae.
The second major division is best commenced with P. niteschi, from a
widespread caprimulgid (Caprimulgiformes). It is the only species to retain
a fully developed tritosternum. However, from this point onward, it becomes
increasingly difficult to correlate morphology and host-specificity. Thus the
three major dichotomies used in the key in Section IV above (those depending
on the degree of posterior truncation of the podonotal shield, the presence
or absence of the postanal seta, and the condition of the pygidial shield)
all lead to incongruities with the system based on host order that will now
largely be followed.
Both P. echinatus, an isolated and widespread species from hirundinids,
and some species from meliphagids (v. infra) show accessory podonotal
shields, but the two groups do not seem otherwise related. Likewise, P. pittae*
(from an Australian pittid) is not necessarily close to the widespread sylviid
parasite P. acrocephali (v. infra) merely because both have a pair of enlarged
setae set close together on the posterior margin of the podonotal shield. It
is possibly closer to P. cractici,* P. motacillae, and P. langei, a trio comprising
members of the so-called sairae species-group of Fain (1959, 1962c). Of these,
the first is a parasite of three genera of Cracticidae (a peculiarly Australian
family), the second is widespread and recorded from a variety of hosts, while
the third is known only from the original record from a hirundinid in the
U:S:S:R:
There now follows a group of eleven species from the Muscicapidae (or
families sometimes included therein, see Keast, 1961). As mentioned above,
it is difficult to arrange them naturally, but the degree of truncation of the
podonotal shield has been given precedence over the absence of the postanal
seta, and the division of the pygidial shield has been considered least likely
to indicate true relationships. These criteria yield the sequences (only non-
muscicapine hosts are indicated)—(i) species with podonotal shield not
truncate, pygidial shield entire, and postanal seta present: P. orthonychus*
(from a timaliid), P. microecae,* P. terpsiphonei, P. bradypteri (from a
sylviid), and P. acrocephali (again from a sylviid) (P. bradypteri is the
only species of the entire sequence to lack peritremes, but this alone does
not, I believe, necessarily make it more closely related to any of the species
listed above which also lack this character, i.e. the other species formerly
included in Passeronyssus and Cas, see synonymy of Ptilonyssus in Section
TV above. Clearly, many characters are polyphyletic in this weakly sclerotized
ROBERT DOMROW 399
group of endoparasites. P. bradypteri is therefore placed before P. acrocephali,
the first species of the sequence to show an anterodorsal process on coxa II.
This latter character then persists throughout the sequence, except for the
last species, P. psophodae); (i%) similar to (7), but postanal seta absent:
P. monarchae ;* (iii) similar to (7), but podonotal shield truncate: P. setosae*
and P. gerygonae* (from a sylviid) ; (iw) similar to (iii), but pygidial shield
divided: P. rhipidurae ;* and (v) similar to (iii), but postanal seta absent:
P. macclurei and P. psophodae* (from a faleunculid).
Of these eleven species, four have been recorded outside Australia.
P. terpsiphonei has a very wide host range in Africa and S. E. Asia.
P. bradypteri and P. acrocephali were originally recorded from Africa and
Kurope, respectively (the Australian host of the latter is also a species
of Acrocephalus Naumann). P. macclurei was described from a Malayan
species of Rhipidwra Vigors and Horsfield, and is restricted, in Australia, to
R. leucophrys. Of the other three Australian species of Rhipidura, R. fuli-
ginosa carries only P. rhipidurae, while the two rain-forest species, R. setosa
and Rk. rufifrons, each bear a variant of a third species, P. setosae. P.
monarchae is known only from the original record, but P. microecae is known
from two species of Microeca Gould, and P. orthonychus and P. gerygonae
have both been taken a second time in their original host species. P. psophodae
is certainly restricted to Psophodes olivaceus.
The family Muscicapidae is cosmopolitan, and, although it has radiated
considerably in Australia, the parasites of muscicapines, at least, may be
expected outside Australia, just as the widespread P. terpsiphonei is estab-
lished in a wide range of Australian genera. The parasites of such peculiarly
Australian branches of this radiation as the pachycephalines and malurines,
however, may be restricted to this region. Finally, this group of eleven
species from muscicapids provides the best Australian example of host-
specificity at three levels: its members may be restricted to a single host
family, genus, or species.
P. grallinae* is an isolated species restricted to Grallina cyanoleuca,
the sole Australian member of the small and aberrant Australopapuan family
Grallinidae (see also diagnosis for P. struthideae in Section IV above).
There are four families of Old World and Australian nectar-feeders, the
Zosteropidae (silvereyes), the Dicaeidae (flower peckers), the Nectariniidae
(sunbirds), and the Meliphagidae (honeyeaters). P. ruandae and P. dicaei,*
two species with short, characteristically broad podonotal shields, are
restricted to the first two, and P. cinnyris to the third of these families.
P. dicaei is still only known from the original record from Dicaeum Cuvier
in Australia, but the other two species are common in other members
of their respective families both in S. E. Asia and Africa.
Some evidence is given above for a burst of speciation within Ptilonyssus
in peculiarly Australian flycatchers. A similar burst is perhaps still in
progress among the parasites of honeyeaters (which, unlike the cosmopolitan
flycatchers, are virtually restricted to Australia), and variants of both
P. philemoni and P. gliciphilae have been described in Section IV above.
Such morphological gaps, correlated with host-specificity, are suggestive of
isolation at a subspecific level, but it is difficult to confirm sexual isolation
in a group with such specialized habits. In any case, mere experimental
confirmation of sexual isolation is of doubtful value, and since it is as much
an error to describe a true species as a subspecies as the reverse, the use
of trinomina for these variants has been foregone (Mayr et al., 1953).
Only one of the three major dichotomies discussed above under the species
from flycatchers is available to divide the species from honeyeaters, viz.
400 THE NASAL MITES OF QUEENSLAND BIRDS
the degree of truncation of the podonotal shield (the pygidial shield is
variable in many species, and the postanal seta is always present). The
application of this criterion yields the following two sequences—(i) podonotal
Shield not truncate: P. myzanthae,* P. philemoni,* P. balimoensis, P.
myzomelae,* and P. gliciphilae;* and (ti) podonotal shield truncate:
P. lymozemae,* P. stomioperae,* P. thymanzae,* and P. meliphagae.* In
addition to this posterior truncation, there is also a lateral reduction of the
podonotal shield in some species, leaving free: accessory shieldlets. This
occurs in both sequences, the species concerned being P. gliciphilae, P.
thymanzae, and P. meliphagae (see note on P. echinatus above). As the
honeyeaters are often gregarious, and several species may be seen feeding
in the same flowering tree, strict host-specificity is not to be expected. Indeed,
double infestations are not uncommon, see Section VII below. Thus P.
thymanzae is known from eight species belonging to four genera, and
P. philemoni from six species of four genera, though it should be noted that
these genera are all toward the end of the sequence given by Leach (1958).
Most of the remaining species are known from two, or occasionally three
host-genera, but there is evidence that both P. myzomelae and P. lymozemae
and restricted to Myzomela Vigors and Horsfield. The striking P. balimoensis,
originally described from an unidentified meliphagid in New Guinea, occurs
in Meliphaga macleayana in Australia.
The isolated P. nudus keys out near the parasites of muscicapids, but
is restricted to the introduced domestic sparrow (Fringillidae), and, like
many of the species with a complete opisthonotal shield parasitic in other
fringillids and ploceids, retains the metasternal setae.
P. sturnopastoris is a parasite of S. E. Asian and Australian sturnids.
P. sphecotheris* and P. trouessarti are restricted to Australian oriolids,
the former to the two species of Sphecotheres Vieillot and the latter to the
two species of Oriolus Linnaeus. They have characteristically cordate
podonotal shields, but key out separately because the postanal seta is absent
in the former and present in the latter.
The imperfectly known P. novaeguineae was described from a paradiseid
Species which penetrates into Australia.
This leaves P. cerchneis, a characteristic and widespread species
restricted to the genus Falco Linnaeus (Falconiformes). It is perhaps related
to the group of species from neotropical falconiforms detailed by Pereira
and de Castro (1949) and Fain and Johnston (1966).
(h) Sternostoma
The members of this nondescript and difficult genus, unlike the preceding
one (Ptilonyssus), show no predilection for passeriform hosts, and host-
specificity is normally above the familial level. S. thienponti is common in
Australian cracticids, although the only previous records are from African
and Asian dicrurids. This finding is of interest, as Thomson (1964) notes
that the relationships of the dicrurids are a “matter of speculation”. The
following four species are virtually cosmopolitan, or at least widespread in
the Old World: 8. cooremani in two coraciiform families, S. boydi in various
charadriiforms, and S. dureni and S. technawi in turdids.
S. cuculorum has numerous hosts, among which cuculids and muscicapids
figure prominently. Of the three recorded cuculid hosts, Cacomantis vario-
losus usually chooses birds that build open, cup-shaped nests in small trees
etc., particularly muscicapids, as foster-parents, and Cayley (1963) notes
that the remaining three Australian hosts also build cup-shaped nests.
ROBPRT DOMROW 401
O. pyrrhophanus, on the other hand, prefers hosts which build a domed nest
near the ground, while the larger Hudynamys orientalis prefers the nests
of orioles and large meliphagids. If I have not over-simplified my concept
of S. cuculorum, these findings differ from those of Strandtmann and Furman
(1956) and Clark (1968), who found that, in the parasitic American cowbirds
(Icteridae, Passeriformes), “host specificity of nasal mites is operative even
in the face of ample opportunities for cross transmission of parasites”.
S. tracheacolum, a pest in aviaries, particularly of fringillids and ploceids,
tends to spread to unusual hosts in these circumstances. It has also been
recorded from a wide range of small passeriforms in the wild (Fain and
Hyland, 1962a). S. paddae, known only from the original record, is associated
with conjunctivitis in an Oriental ploceid, and S. eryptorhynchum is wide-
spread in the house-sparrow.
The remaining three species (S. gliciphilae,* S. neosittae,* and S. fulicae)
have not been collected since the original series. The first is the only species
of the genus recorded from a peculiarly Australian host family (Melipha-
gidae), but the second may be expected to occur in other sittids overseas.
The third is a parasite of the coot (Rallidae).
(1) Boydaia
This large genus, unlike the remaining three speleognathine genera
discussed below, is essentially restricted to passeriform hosts, although, of
the Australian species, B. falconis is widespread in the genus Falco. B.
spatulata and B. crassipes are both widespread parasites of a variety of
hosts, but B. sturni, B. zosteropis, and B. hirundoae, though widespread,
are restricted to sturnids, zosteropids, and hirundinids, respectively. B.
myzomelae* and B. maluri* are newly described from two peculiarly
Australian groups, the Meliphagidae and Malurinae (Sylviidae).
(j) Neoboydaia
The five Australian members of this genus (NV. philomachi, N. psittaculae,
N. aureliani, N. merops, and N. colymbiformi) are all widespread, showing
a low level of host-specificity for five different non-passeriform orders
(Charadrii-, Psittaci-, Strigi-, Coracii-, and Podicipiformes, respectively).
(k) Ophthalmognathus
O. schoutedeni is widespread in ardeids, and O. striatus in columbids.
O. accipitris,* newly described from an Australian accipitrid, may also have
a wider range. ‘
(1) Speleognathopsis
S. benoiti is a widespread parasite of various charadriiforms, while
S. galli and S. porphyrionis* are known, as yet, only from galliforms in
Africa and a gruiform in Australia, respectively.
(m) Turbinoptes
T. strandtmanni is a widespread parasite of various charadriiforms.
(n) Passerrhinoptes
P. pomatostomi* is known only from an Australian timaliid, but may
extend, with this family, as far as Africa.
(0) Oxleya*
O. podargi* was described from an Australian caprimulgiform, which.
_ however, extends into the Orient.
402 THE NASAL MITES OF QUEENSLAND BIRDS
(p) Schoutedenocoptes
S. numidae is known only from a domesticated numidid.
(q) Rhinoptes
R. gallinae was also described frém a domesticated phasianid, but also
occurs wild in an Australian member of the same host-family.
(r) Mycteroptes
M. basilewskyi is widespread in coraciids.
.In conclusion, the anatomy oi all birds is very similar, and their
relatively recent emergence, their low rate of extinction, and their poor
fossil record have further contributed to the difficulty in providing a classi-
fication and a detailed history of the phylogeny ef the various bird orders.
The birds began to diverge from reptilian stock in the late Jurassic
(150 million years ago), shortly after the emergence of the first mammals,
but the earliest line (the Archaeornithes, which included only Archaeopteryx
Meyer) is long extinct.
The Neornithes, which include the remaining fossil groups as well as
all living birds, arose in the Cretaceous (120 m. years ago), and even by
this time some birds had evolved which can be referred to orders still extant.
During the Tertiary, the birds (and the mammals) gained ascendancy over
the reptiles, and most of the orders, both of land-and water-birds, evolved
(Eocene, 50 m. years ago), modern genera began to appear (Oligocene-Miocene,
35-25 m. years ago), and many modern species were already present
(Pliocene, 10 m. years ago).
Both groups of modern birds are cosmopolitan. The palaeognaths (or
flightless ratites) include only the ostrich of Africa and (formerly) S. W.
Asia, the emu and cassowaries of Australia and New Guinea, the kiwis and
recently extinct moas of New Zealand, the rheas of South America, etc. To
the best of my knowledge, the only ratites examined for nasal mites are
three emus (Dromaius Vieillot) (v. infra) and two cassowaries (Casuarius
Brisson) (Wilson, personal communication). All were uninfested. The
neognaths therefore include all the known hosts of nasal mites. They
comprise all the remaining orders, but, as noted above, it is difficult to
classify a group so diverse as to include the tinamous, penguins, grebes,
petrels, ducks, hawks, pigeons, parrots, owls, swifts, etc. on the one hand, not
to mention the equally large series of passeriform families on the- other
(Young, 1950; Thomson, 1964).
It seems clear, therefore, that mites of three suborders entered the nasal
cavities of birds at an early period, and spread with their hosts, so that
such widely differing groups of birds as grebes, herons, swallows, larks, etc.
now bear their own distinctive mites throughout their cosmopolitan ranges.
The same is true of groups confined to the Old World, even in the case of
families possessing numerous species in Africa and Asia that are represented
in Australia only by one. The Coraciidae, Meropidae, Nectariniidae,
Sturnidae, and Dicruridae come immediately to mind, and all have carried
‘their specific parasite(s) into Australia. A similar situation occurs even
with such introduced species as the starling and house-sparrow, and this
conservatism is opposed only by the parasites of groups of birds, like the
meliphagids, that have been free to radiate in isolation at the end of this
chain in Australia.
VI. PHYLOGENY OF THE RHINONYSSINE GENERA
Although no firm relationship with other arachnids can be established
from the meagre fossil record, mites have been reported from the Devonian
ROBERT DOMROW 405
and Carboniferous, while, by the Tertiary, all the principal families are
represented, particularly in amber (Woolley, 1961). Vitzthum’s concept
(1935, 1942) of the Dermanyssidae (then included in the Laelapidae) as
primitively humus-dwelling predators, various lines of which became para
phages of insects, nidophiles, ectoparasites, and finally endoparasites of a
wide range of vertebrate hosts, was maintained by Zumpt and Patterson
(1951), and Evans and Till (1966) have recently retreated the matter in
detail.*
It therefore seems reasonable to look for the primitive hosts of the
Rhinonyssinae among the land-birds. The simplest species of Tinaminyssus
(those retaining two well developed dorsal shields) are, indeed, those from
pigeons, hawks, and parrots, the one exception, 7’. belopolskii, being a parasite
of herons. The tritosternum, another structure quickly lost in an endo-
parasitic group, is also retained in some species from parrots. Some
fragmentation, or occasionally the disappearance of the opisthonotal shield
takes place in other species from hawks and parrots, and the species from
tinamous (neotropical ground-birds) also show a divided opisthonotal shield.
The species of Tinaminyssus from kingfishers generally lack the opisthonotal
shield, as do those of the related genera Larinyssus and Rallinyssus, T.
tanysipterae (Wilson, 1966a), n. comb., and R. amaurornis Wilson (1965)
being exceptions. Larinyssus is restricted to various waders and Rallinyssus
to rails. The latter, because of its peculiarly displaced stigmata and anal
accoutrements, is presumably the more evolved. The fragmentation of the
dorsal shield in these two genera extends to the podonotal shield, and this
is to be correlated with the decreasing danger of desiccation and anoxia in
the nasal passages of fresh-water and marine birds. The genus Rhinonyssus,
in which the peritremes (porous anterior extensions of the respiratory
stigmata) are lost, closes this line of evolution. The simpler species of this
genus are parasites of penguins and a wide variety of waders, but one
compact species-group with peculiarly modified palpi is found both in ducks
and grebes (Strandtmann, 1959). It should be noted, however, that penguins
are much closer to the Procellariiformes (petrels, albatrosses, etc.) than to
the waders, and that the grebes are not ducklings, as T have heard bushmen
heatedly affirm. In this case, the gregarious habits of the hosts in salt- and
fresh-water are more important in determining host-specificity than true
phylogenetic relationships (Strandtmann, 1958). To recapitulate, in this
Series of four small genera, all with unmodified chelicerae, the movement
has been towards, and not away from water-birds.
A second line of development is to be seen in the two small genera
Ruandanyssus and Rhinoecius, in which the fixed cheliceral digit has been
lost, leaving a faleate movable digit of moderate length. The former
parasitizes a range of passeriform birds, while the latter is restricted to owls.
A third line is characterized by a considerable reduction in size of the
cheliceral digits, with a concomitant attenuation of the distal half of the
cheliceral shafts (the seemingly bulbous, but normally developed basal portion
is drawn out ventrally to form this slender portion). The species of
Ptilonyssus which have retained two large dorsal shields and (often)
*The classification of the two other groups of nasal mites of birds is still fluid.
The Ereynetidae are divided by Fain (1962d) into the Ereynetinae (essentially free-
living predators, but including one ectoparasite of pulmonate gastropods and one
intranasal parasite of a Malaysian bird, see Fain, 1964e, and Fain and Nadchatram,
1962), the Lawrencarinae (intranasal parasites of frogs), and the Speleognathinae
(intranasal parasites of warm-blooded vertebrates). The turbinoptines are close
relatives of epidermoptine genera which scavenge on the skin débris of birds.
404 THE NASAL MITES OF QUEENSLAND BIRDS
metasternal setae are principally parasites of small granivores, while the
remainder have radiated, and are still radiating among a wide range of
passeriforms. An occassional species of this very large genus has been
recorded from non-passeriforms (hawks, swifts, nightjars, and bee-eaters),
but none is known from water-birds. ‘The species of Sternostoma, found as
a rule deeper in the nasal tract, have lost their peritremes, and one, with
a reduced opisthonotal shield, is found in waders. The remainder are
parasites particularly of passeriforms, but, rather more frequently than is
the case with Ptilonyssus, also of non-passeriform groups.
‘These three lines are the subfamilies Rhinonyssinae, Rhinoeciinae, and
Ptilonyssinae discussed, and abandoned in Section IV above, together with
Bregetova’s theory (1964) of a diphyletic origin of the rhinonyssines. Indeed,
the rhinonyssines with a well developed tritosternum are so little differentiated
from some of the ectoparasites of mammals and birds, e.g. Ornithonyssus
Sambon and Pellonyssus Clark and Yunker, that Zumpt and Till (1961)
placed Ruandanyssus and Astridiella (here considered a synonym of
Ptilonyssus) among the macronyssine rather than the rhinonyssine genera.
If a distinction must be maintained between the Macronyssinae and the
Rhinonyssinae, it can be made only on ecological grounds, because the
presence or absence of the tritosternum is useless as a taxonomic character,
as discussed above in the synonymies of Tinaminyssus, Rhinoecius, and
Ptilonyssus. Indeed, even this ecological separation is weakened by the habits
of some related species. The tropical fowl-mite, Ornithonyssus bursa (Berlese)
(Macronyssinae), unlike the northern fowl-mite, O. sylviarum (Canestrini
and Fanzago), does not spend its whole time on its host (Furman, 1963),
and the chicken-mite, Dermanyssus gallinae (Degeer) (Dermanyssinae) feeds
only for one or two hours at night, spending the rest of the day in crevices
in fowlhouses (Kirkwood, 1963). Such behaviour undoubtedly led to the
entry of “protorhinonyssines” into the intricate, warm, and moist nasal
passages of birds. I therefore believe the Rhinonyssinae to be a monophyletic
unit clearly derived, and separable only with difficulty from the Mac-
ronyssinae, the former name having priority.
VII. Host-parasite List
As the revision of the official checklist of Australian birds (Leach
et al., 1926) has not yet been completed, I have used the classification of
Leach (1958)* in the present study. This authority lists 735 species, 330
genera, 82 families, and 20 orders (including 11 genera and 17 species of
introduced and vagrant birds), while an unpublished checklist of Queensland
birds on a similar basis (Lavery, personal communication) lists 570 species
and 280 genera. It is therefore clear that, since Queensland, of all the states,
has by far the largest proportion of the birds on the Commonwealth list
(78% of the species, and 85% of the genera), an intensive study of the fauna
of that state will have relevance beyond its borders. Leach’s system is
summarized in Tables 2 and 3, with an indication of the fauna still to be
examined. However, as with many of Australia’s rarer marsupials, it is
unlikely that some groups, e.g. night parrots (Geopsittacus Gould) and
scrub-birds (Atrichornis Stejneger), will ever be thoroughly examined for
parasites.
*As I am not a bird taxonomist, I have accepted uncritically the Latin names of
this authority, although several are obviously in error. Thus, while the use of
adjectival specific names with masculine terminations (pyrrhopygius, sanctus) in the
feminine genus Halcyon contravenes Art. 30, the original spelling of the specific name
tenuirostris, not being adjectival, stands under Art. 32, and cannot be forced into a
neuter form on its transferal to Edoliisoma Pucheran. I have, however, dispensed with
the hyphen in specific names such as novaehollandiae (Art. 32).
ROBERT DOMROW 405
I have examined just over half (287 species) of the Queensland total,
while another 43 widespread species known to be infested overseas have not
yet been so found in Australia (some of this latter group occur in Queensland;
others do not). Of those I have examined, 158 species (55%) were infested
(of which eight are likely to yield still further mite species), while a further
51 species can be expected to yield nasal mites. In total, 330 species are
known to have been examined, of which nasal mites have been recorded from
209 (68%).
TABLE 2
Analysis of Australian and Queensland genera and species of birds examined and found infested with
nasal mites
Number of Genera Number of Species
Order —-_— a — _—$—
Aust- Queens- Ex- In- Aust- Queens- Ex- In-
ralia land* amined _ fested ralia land* amined fested
Casuariiformes Lb 2; 2 1 0 2 2 1 0
Galliformes st 8 6 5 3 9 7 5 3
Turniciformes By: 2 2 1 0 8 7 1 0
Columbiformes sf 17 14 12 12 27 22 17 16
Gruiformes .. 12 10 9 7 16 15 9 7
Podicipiformes - 1 1 1 1 3 3 1 1
Sphenisciformes .. 3 1 1 1 5 1 1 1
Procellariiformes .. 16 12 0 0 37 26 0 0
Charadriiformes Ph 38 32 28 20 76 75 44 31
Ciconiiformes of 14 12 11 5 20 21 17 7
Anseriformes — a 15 12 8 4 22 18 11 7
Pelecaniformes as 6 6 4 0 15 14 6 0
Falconiformes a 14 14 8 3 26 24 12 4
Strigiformes By 2 2 2 2 9 8 + 2
Psittaciformes = 23 19 13 12 58 36 20 14
Caprimulgiformes .. 4 4 3 2 8 7 3 2
Coraciiformes uf 7 a 5 4 12 12 10 8
Apodiformes of 3 3 0 0 3 5 0 0
Cuculiformes ue 7 7 6 2 14 12 8 3
Passeriformes so ete 114 93 72 365 255 160 103
Total.. x. se aed) 280 211 150 735 570 330 209
Percentage : -
Australia ae PloOo 85 64 45 100 78 45 28
Queensland .. — 100 75 53 — 100 58 36
Examined ae — — 100 Al — — 100 62
nS Lavery’s figures (personal communication) for the Galliformes, Turniciformes, Gruiformes,
and Charadriiformes have been adjusted to conform to Leach’s order (1958) to facilitate
comparison of the two sets of figures.
The number of individual host-parasite records, however, is somewhat
larger. My own collections have yielded 205 such records (Rhinonyssinae
from 175, Speleognathinae from 23, and Turbinoptinae from 7 hosts), and,
taking into account birds on the Australian list that are known to be infested
Overseas, this number may be increased to 295 (Rhinonyssinae from 248,
Speleognathinae from 37, and Turbinoptinae from 10 hosts). These figures
are neither maximal nor minimal, since some birds examined in small
numbers only will probably prove later to be infested, while others, of which
only a single specimen has been examined, have proved infested (with, in
at least one case, Oreocincla lunulata, two species of mites).
The reason for this difference is that one host-species may harbour two
or more species of mites, and, in fact, 30 hosts are known, in Australia, to
harbour two, seven three, and one four parasites. However, no multiple
infestations were noted in the present study, but the following 22 cases of
natural double infestations occurred, 14 involving two rhinonyssine species,
406
THE NASAL MITES OF QUEENSLAND BIRDS
TABLE 3
Synopsis of genera and species of Australian birds examined and found infested with nasal mites
Order
Family
Casuariiformes
Galliformes
Turniciformes
Columbiformes
Gruiformes
Podicipiformes
Sphenisciformes
Procellariiformes
Charadriiformes
Ciconiiformes
Anseriformes
Pelecaniformes
Falconiformes
Strigiformes
Psittaciformes
Caprimulgiformes
Coraciiformes
_ Apodiformes
Cuculiformes
Passeriformes
Dromaiidae
Casuariidae
Megapodiidae
Phasianidae
Numididae
Turnicidae
Pedionomidae
Columbidae
Rallidae
Megalornithidae
Podicipidae
Spheniscidae
Hydrobatidae
Procellariidae
Pelecanoididae
Diomedeidae
Laridae
Stercorariidae
Haematopodidae
Charadriidae
Recurvirostridae
Scolopacidae
Rostratulidae
Jacanidae
Glareolidae
Burhinidae
Otitidae
Threskiornithidae
Ciconiidae
Ardeidae
Anatidae
Phalacrocoracidae
Anhingidae
Sulidae
Fregatidae
Phaethontidae
Pelecanidae
Accipitridae
Falconidae
Pandionidae
Strigidae
Tytonidae
Psittacidae
Podargidae
Aegothelidae
Caprimulgidae
Coraciidae
Alcedinidae
Meropidae
Apodidae
Cuculidae
Menuridae
Pittidae
Atrichornithidae
Hirundinidae
Muscicapidae
Campophagidae
Timaliidae
Sylviidae
Total
—
OVS met OS me ND OD me mm mm 0 0D me OD OD BD OO OF OO ee ee eT RO
— pt —)
—
bo
—
ix)
Number of Genera
Ex- In-
amined fested
—_
SCMNDK ORCC OC HR KH ONOK RK NWO F
—
SOOnocoCaArFOOCOCROCOCOCOFrFOCANOCOOFNOOS
—
CORD RH 09 OH SO St et et et ett OO dt OS et 00 de
—
= —
ONWNWOCONCRKFONORE NKR RK Of NR RK OF NCCOCOCOCCrR AS
Pm UR SD CD eT ND tt eet et et DD me et et et ee
bo bo hm bo
OS G2 BS OS me ad = PO OU OU ee dO
So =
bo
ou om
—
—
(Se)
OCAIMNWNHRWHKH OK WE PDA — 10 bd
a
OO
bo
Number of Species
Ex- In-
Total amined fested
i
—
SON eKHSOSCHACCCCH KH OAWGOCOrFNOOCS
1
0
2
2
1
1
0
7
8
1
1
1
0
0
0
0
8
0
1
10
2
4 18 1
1 0 -
1 1
2 2
2 1
1 1
5 5
1 1
4 11
2 1]
5 ye 8
1 1
4 1
2 0
0
bo
=
NPWNWOONORKF OCOOOCOK OR HK RK REP ONN OW Oe
DNWOROHOWOH OP ROP BHEHONNOSOOOOCONAICOCOCON
=
ROBERT DOMROW 407
Tasie 3—Continued
Synopsis of genera and species of Australian birds examined and found infested with nasal
mites—Continued
Number of Genera Number of Species
Order Family a
Ex- In- Ex- In-
Total amined fested Total amined fested
Passeriformes Artamidae
Grallinidae
Pachycephalidae
Falcunculidae -
Turdidae
Pycnonotidae
Cracticidae
Paridae
Sittidae
Certhiidae
Zosteropidae
Dicaeidae
Nectariniidae
Meliphagidae 2
Motacillidae
Alaudidae
Fringillidae
Ploceidae 1
Sturnidae
Oriolidae
Dicruridae
Paradiseidae
Ptilonorhynchidae
Corvidae
—
COs. D>
—
—
BO DS m= = DO GO OU Re DD ee ee OOOO NY
>
to
ROWE PON ORK NOH Oe OOO RmOP MOO oF
_—
CO Rm eb WO Re Oe Ne ee OW CONN ee
bo
AOCRK PWOWANNOK ORCA
—
—)
DO DD = = ROO OL OL DO ee eR Re OO OWN Oe &
Sa td I a a So ed dn
—_
and eight one rhinonyssine and one speleognathine species (turbinoptines
were never involved): Tinaminyssus myristiciwworae and T. welchi from
Myristicwora spilorrhoa; Rhinonyssus minutus and Speleognathopsis benoiti
from Charadrius melanops (twice); Ruandanyssus terpsiphonei and
Ptilonyssus struthideae from Struthidea cinerea; Ruandanyssus terpsiphonei
and Sternostoma cuculorum from Myiagra rubecula; Ptilonyssus triscutatus
and Neoboydaia merops from Merops ornatus; Ptilonyssus dioptrornis and
Sternostoma cuculorum from Eopsaltria capito; Ptilonyssus cractici and
Sternostoma thienponti from Gymnorhina tibicen; Ptilonyssus terpsiphonei
and Sternostoma cuculorum from Arses kaupi; Ptilonyssus monarchae and
Sternostoma cuculorum from Monarcha trivirgata; Ptilonyssus ruandae and
Sternostoma zosteropis from Zosterops lateralis; Ptilonyssus myzanthae and
P. thymanzae from Myzantha melanocephala (four times); Ptilonyssus
myzanthae and P. thymanzae from Myzantha flavigula; Ptilonyssus philemoni
and Boydaia spatulata from Philemon corniculatus; Ptilonyssus gliciphilac
and Boydaia spatulata from Myzomela pectoralis ; Ptilonyssus lymozemae and
Boydaia myzomelae from Myzomela sanguinolenta; Ptilonyssus cerchneis and
Boydaia falconis from Falco berigora (twice) ; and Sternostoma dureni and
S. technawi from Oreocincla lunulata.
The complete host-parasite list occupies Table 4 below, in which the
following typographical conventions have been used: (i) asterisks indicate
birds known to occur in Australia as introductions or vagrants, but not
listed in Leach (1958); (i) scientific names of birds printed in roman
rather than in italics indicate species not examined in the present survey:
and (ti) scientific names of mites similarly printed indicate host-parasite
relationships established overseas, but not yet noted in Australia.
408
TABLE 4
THE NASAL MITES OF QUEENSLAND BIRDS
List of Australian birds examined and their recorded nasal mite parasites
Dromaius novaehollandiae
(Latham)
Megapodius freycinet Gaimard
Aleectura lathami Gray
Gallus gallus (Linnaeus)*
Synoicus australis (Latham)
Numida meleagris (Linnaeus)*
Turnix varia (Latham)
Piilinopus regina Swainson
Piilinopus superbus (Temminck)
Megaloprepia magnifica (Tem-
minck)
Myristicivora spilorrhoa (Gray)
Columba livia Gmelin*
Columba norfolciensis Latham
Macropygia phasianella (Tem-
minck)
Streptopelia chinensis (Scopoli)
Streptopelia senegalensis (Lin-
naeus)
Geopelia placida Gould
Geopelia cuneata (Latham)
Geopelia humeralis (Temminck)
Chalcophaps chrysochlora (Wagler)
Phaps chalcoptera (Latham)
Geophaps scripta (Temminck)
Ocyphaps lophotes (Temminck)
Leucosarcia melanoleuca (Latham)
Rallus pectoralis Temminck
Hypotaenidia philippensis (Lin-
naeus)
Porzana tabuensis (Gmelin)
Poliolimnas cinereus (Vieillot)
‘Amaurornis ruficrissus (Gould)
Gallinula tenebrosa Gould
Porphyrio melanotus Temminck
Fulica atra Linnaeus
Grus rubicundus (Perry)
Casuariiformes
Dronsaiidae
Emu
Galliformes
Megapodiidae
Serub-fowl
Brush-turkey
Phasianidae
Domestic fowl
Brown quail
Numididae
Guinea-fowl
Turniciformes
Turnicidae
Painted quail
Columbiformes
Columbidae
Red-crowned pigeon
Purple-crowned pigeon
Wompoo pigeon
Torres Strait pigeon
Domestic pigeon
White-headed pigeon
Brown pigeon
‘Indian turtle-dove
Senegal dove
Peaceful dove
Diamond-dove
Bar-shouldered dove
Green-winged pigeon
Common bronzewing
Squatter-pigeon
Crested pigeon
Wonga pigeon
Gruiformes
Rallidae
Lewin water-rail
Banded landrail
Spotless crake
White-browed crake
Bushhen
Dusky moorhen
Eastern swamphen
Coot
Megalornithidae
Broiga
Speleognathopsis galli
Rhinoptes gallinae
Rhinoptes gallinae
Speleognathopsis galli
Schoutedenocoptes numidae
Tinaminyssus
Tinaminyssus
Tinaminyssus
Tinaminyssus
Tinaminyssus
Tinaminyssus
ptilinopi
megaloprepiae
myristicivorae
welchi
mellow
columbae
Ophthalmognathus striatus
Tinaminyssus
Tinaminyssus
Tinaminyssus
Tinaminyssus
Tinaminyssus
Tinaminyssus
Tinaminyssus
Tinaminyssus
Tinaminyssus
Tinaminyssus
Tinaminyssus
Tinaminyssus
Tinaminyssus
mellot
macropygiae
mellow
melloi
geopeliae
geopeliae
geopeliae
hirius
phabus
ocyphabus
ocyphabus
columbae
mello
Rallinyssus gallinulae
Rallinyssus gallinulae
Rallinyssus congolensis
Rallinyssus amaurornis
Rallinyssus caudistigmus
Rallinyssus gallinulae
Speleognathopsis porphyrionis
Rallinyssus caudistigmus
Sternostoma fulicae
ROBERT DOMROW
TABLE 4—Continued
409
List of Australian birds examined and their recorded nasal mite parasites—Continued
Podiceps ruficollis (Vroeg)
Pygoscelis adeliae (Hombron and
Jacquinot)*
Chlidonias hybrida (Pallas)
Chlidonias Jleucoptera (Tem-
minck) F
Gelochelidon nilotica (Gmelin)
Sterna hirundo Linnaeus
Sterna bergit Lichtenstein
Sterna fuscata Linnaeus
Larus novaehollandiae Stephens
Larus dominicanus Lichtenstein*
Haematopus ostralegus Linnaeus
Arenaria interpres (Linnaeus)
Erythrogonys cinctus Gould ~
Lobibyx novaehollandiae
(Stephens)
Lobibyx miles (Boddaert)
Zonifer tricolor (Vieillot)
Squatarola squatarola
naeus)
Pluvialis dominicus (Miiller)
(Lin-
Charadrius alexandrinus Linnaeus
Charadrius melanops Vieillot
Charadrius hiaticula Linnaeus*
Himantopus leucocephalus Gould
Recurvirostra novaehollandiae
Vieillot
Numenius madagascarensis (Lin-
naeus)
Numenius phaeopus (Linnaeus)
Mesoscolopax minutus (Gould)
Limosa lapponica (Linnaeus)
Limosa limosa (Linnaeus)
Tringa hypoleueca Linnaeus
Tringa brevipes (Vieillot)
Tringa glareola Linnaeus
Tringa nebularia (Gunnerus)
Podicipiformes
Podicipidae
Little grebe
Sphenisciformes
Spheniscidae
Adélie penguin
Charadriiformes
Laridae
Whiskered tern
White-winged black tern
Gull-billed tern
Common tern
Crested tern
Sooty tern
Silver gull
Southern black-backed gull
Haematopodidae
Pied oystercatcher
Charadriidae
Turnstone
Red-kneed dotterel
Spur-winged plover
Masked plover
Banded plover
Grey plover
Eastern golden plover
Red-capped dotterel
Black-fronted dotterel
Ringed plover
Recurvirostridae
White-headed stilt
Red-necked avocet
Scolopacidae
Eastern curlew
Whimbrel
Little whimbrel
Bar-tailed godwit
Black-tailed godwit
Common sandpiper
Grey-tailed tattler
Wood-sandpiper
_ Greenshank
Rhinonyssus poliocephali
Neoboydaia colymbiformi
Rhinonyssus sphenisci
Larinyssus orbicularis
Sternostoma boydi
Larinyssus orbicularis
Larinyssus orbicularis
Larinyssus orbicularis
Larinyssus orbicularis
Sternostoma boydi
Turbinoptes strandtmanni
Larinyssus orbicularis
Rhinonyssus coniventris
Sternostoma boydi
Rhinonyssus himantopus
Speleognathopsis benoiti
Rhinonyssus himantopus
Rhinonussus himantopus
Speleognathopsis benoiti
Rhinonyssus coniventris
Speleognathopsis benoiti
Rhinonyssus minutus
Rhinonyssus coniventris
Rhinonyssus minutus
Rhinonyssus coniventris
Rhinonyssus himantopus
Speleognathopsis benoiti
Rhinonyssus minutus
Rhinonyssus coniventris
Rhinonyssus himantopus
Turbinoptes strandimanni
Turbinoptes strandtmanni
Neoboydaia philomachi
Sternostoma boydi
Turbinoptes strandtmanni
Rhinonyssus coniventris
Sternostoma boydi
Neoboydaia philomachi
Rhinonyssus rhinolethrum
Sternostoma boydi
41:0
TABLE 4—Continued
THE NASAL MITES OF QUEENSLAND BIRDS
List of Australian birds examined and their recorded nasal mite parasites—Continued
Crocethia alba (Vroeg)
Erolia ruficollis (Pallas)
Erolia acuminata (Horsfield)
Erolia melanotos (Vieillot)
Erotia ferruginea (Brunnich)
Erolia alpina (Linnaeus)*
Calidris canutus (Linnaeus)
Gallinago hardwickw (Gray)
Philomachus pugnax (Linnaeus)*
Trediparra gallinacea (Temminck)
Stiltia isabella (Vieillot)
Glareola pratincola (Linnaeus)
Burhinus magnirostris (Latham)
Hupodotis australis (Gray)
Threskiornis molucca (Cuvier)
Threskiornis spinicollis (Jameson)
Plegadis falcinellus (Linnaeus)
Platalea regia Gould
Platalea flavipes Gould
Xenorhynchus asiaticus (Latham) —
Egretta intermedia (Wagler)
Egretta alba (Linnaeus)
Egretta garzetta (Linnaeus)
Bubulcus ibis (Linnaeus)*
Notophoyx novaehollandiae
(Latham)
Notophoyx pacifica (Latham)
Notophoyx picata (Gould)
Demigretta sacra (Gmelin)
Nycticorax caledonicus (Gmelin)
Butorides striata (Linnaeus)
Ixobrychus minutus (Linnaeus)
Anser anser (Linnaeus)*
_Anseranas semipalmata (Latham)
Nettapus pulchellus Gould
Chenonetta jubata (Latham)
Dendrocygna arcuata (Horsfield)
Tadorna radjah (Garnot)
Anas superciliosa Gmelin
Anas querquedula Linnaeus
Anas gibberifrons (Miller)
Anas platyrhynchos Linnaeus*
Aythya australis (Kyton)
Sanderling
Red-necked stint
Sharp-tailed sandpiper
American pectoral sand-
piper
Curlew-sandpiper
Dunlin
Knot
Australian snipe
Ruff
Jacanidae
Lotus-bird
Glareolidae
Australian pratincole
Oriental pratincole
Burhinidae
Southern stone-curlew
Otitidae
Australian bustard
Ciconiiformes
Threskiornithidae
Australian white ibis
Straw-necked ibis
Glossy ibis
Royal spoonbill
Yellow-billed spoonbill
Ciconiidae
Jabiru
Ardeidae
Plumed egret
White egret
Little egret
Cattle egret
White-faced heron
White-necked heron
Pied heron
Reef-heron
Nankeen night heron
Mangrove-heron
Little bittern
Anseriformes
Anatidae
Domestic goose
Pied goose
Green pigmy goose
Maned goose
Whistling tree-duck
White-headed shelduck
Black duck
Garganey teal
Grey teal
Mallard
Hardhead
Rhinonyssus coniventris
Sternostoma boydi
Rhinonyssus coniventris
Neoboydaia philomachi
Rhinonyssus coniventris
Rhinonyssus coniventris
Sternostoma boydi
Neoboydaia philomachi
Lariyssus benoiti
Larinyssus benoiti
Tinaminyssus belopolskii
Tinaminyssus belopolskii
Tinaminyssus belopolskii
Tinaminyssus belopolski
Tinaminyssus belopolskiz
Ophthalmognathus schoutedeni
Tinaminyssus belopolskii
Rhinonyssus rhinolethrum
Rhinonyssus rhinolethrum
Rhinonyssus rhinolethrum
Rhinonyssus rhinolethrum
Rhinonyssus rhinolethrum
Rhinonyssus rhinolethrum
Rhinonyssus rhinolethrum
ROBERT DOMROW
TABLE 4—Continued
411
List of Australian birds examined and their recorded nasal mite parasites—Continued
Phalacrocorax sulcirostris
(Brandt)
Phalacrocorax varius (Gmelin)
Phalacrocorax melanoleucus
(Vieillot)
Anhinga novaehollandiae (Gould)
Sula dactylatra Lesson
Pelecanus conspicillatus Tem-
minck
Accipiter novaehollandiae (Gmelin)
Accipiter fasciatus (Vigors and
Horsfield)
Accipiter cirrocephalus (Vieillot)
Aquila audax (Latham)
Haliaeetus leucogaster (Gmelin)
Haliastur sphenurus (Vieillot)
Milvus migrans (Boddaert)
Elanus notatus Gould
Aviceda subcristata (Gould)
Falco longipennis Swainson
Falco berigora Vigors and Hors-
field
Falco cenchroides Vigors
Horsfield
and
Ninox novaeseelandiae (Gmelin)
Ninox connivens (Latham)
Tyto alba (Scopoli)
Tyto novaehollandiae (Stephens)
Agapornis roseicollis (Vieillot)*
Trichoglossus moluccanus
(Gmelin)
Trichoglossus chlorolepidotus
(Kuhl)
Psitieuteles versicolor (Lear)
Glossopsitia pusilla (Shaw)
Calyptorhynchus bankstwi (Latham)
Kakatoe galerita (Latham)
Kakatoe roseicapilla (Vieillot)
Kakatoe sanguinea (Gould)
Leptolophus hollandicus (Kerr)
Aprosmictus scapularis (Lich-
tenstein)
Aprosmictus erythropterus
(Gmelin)
Platycercus elegans (Gmelin)
Platycercus adscitus (Latham)
Platycercus eximius (Shaw)
Pelecaniformes
Phalacrocoracidae
Little black cormorant
Pied cormorant
Little pied cormorant
Anhingidae
Australian darter
Sulidae
Masked gannet
Pelecanidae
Australian pelican
Falconiformes
Accipitridae
Grey goshawk
Australian goshawk
Collared sparrow hawk
Wedge-tailed eagle
White-breasted sea-eagle
Whistling eagle
Black. kite
Australian black-shouldered
kite
Crested hawk
Faleconidae
Little falcon
Brown hawk
Nankeen kestrel
Strigiformes
Strigidae
Boobook owl
Winking owl
Tytonidae
Barn-owl
Masked owl
Psittaciformes
Psittacidae
Peach-faced lovebird
Rainbow-lorikeet
Scaly-breasted lorikeet
Varied lorikeet
Little lorikeet
Red-tailed black cockatoo
White cockatoo
Galah
Little corella
Cockatiel
King-parrot
. Red-winged parrot
Crimson rosella
Pale-headed rosella
Eastern rosella
Ophthalmognathus accipitris
Tinaminyssus epileus
Ptilonyssus cerchneis
Boydaia falconis
Ptilonyssus cerchneis
Rhinoecius cooremani
Rhinoecius tytonis
Neoboydaia aureliani
Neoboydaia psittaculae
Tinaminyssus trichoglossi
Neoboydaia psittaculae
Tinaminyssus trichoglosst
Neoboydaia psittaculae
Tinaminyssus trichoglosst
Tinaminyssus kakatuae
Tinaminyssus kakatuae
Tinaminyssus kakatuae
Tinaminyssus aprosmicti
Tinaminyssus aprosmicti
Tinaminyssus aprosmicti
442
TABLE 4—Continued
THE NASAL MITES OF QUEENSLAND BIRDS
Ist of Australian birds examined and their recorded nasal mite parasites—Continued
Barnardius barnardi (Vigors and
Horsfield)
Psephotus varius Clark
Psephotus haematonotus (Gould)
Neophema pulchella (Shaw)
Melopsitiacus undulatus (Shaw)
Podargus strigoides (Latham)
Aegotheles cristata (Shaw)
Caprimulgus macrurus Horsfield
Hurystomus orientalis (Linnaeus)
Alcyone azurea (Latham)
Alcyone pusilla (Temminck)
Dacelo gigas (Boddaert)
Dacelo leachit Vigors and Hors-
field
Halcyon pyrrhopygius Gould
Halcyon sanctus Vigors
Horsfield
Halcyon macleayu Jardine and
Selby
Haleyon chloris (Boddaert)
and
Merops ornatus Latham
Cuculus saturatus Blyth
Cacomantis pyrrhophanus
(Vieillot)
Cacomantis variolosus
and Horsfield)
Chalcites basalis (Horsfield)
Chalciies plagosus (Latham)
Scythrops novaehollandiae Latham
Hudynamys orientalis (Linnaeus)
Centropus phasianinus (Latham)
(Vigors
Pitta versicolor Swainson
Hirundo neoxena Gould
Hirundo rustica Linnaeus*
Ring-neck parrot
Mulga parrot
Red-backed parrot
Turquoise parrot
Budgerygah
Caprimulgiformes
Podargidae
Tawny frogmouth
Aegothelidae
Owlet-nightjar
Caprimulgidae
Large-tailed nightjar
Coraciiformes
Coraciidae
Eastern broad-billed roller
Alcedinidae
Azure kingfisher
Little kingfisher
Laughing kookaburra
Blue-winged kookaburra
Red-backed kingfisher
Sacred kingfisher
Forest kingfisher
Mangrove-kingfisher
Meropidae
'Rainbow-bird
Cuculiformes
Cuculidae
Oriental cuckoo
Fan-tailed cuckoo
Brush cuckoo
Horsfield bronze-cuckoo
Golden bronze-cuckoo
Channel-billed cuckoo
Koel ;
Pheasant-coucal
Passeriformes
Pittidae
Noisy pitta
Hirundinidae
Welcome swallow
Common swallow
Tinaminyssus aprosmicti
Tinaminyssus aprosmicti
Tinaminyssus aprosmicti
Sternostoma tracheacolum
Oxleya podargi
Ptilonyssus nitzschi
Mycteroptes basilewsky2
Tinaminyssus daceloae
Ptilonyssus cractici
Sternostoma cooremant
Tinaminyssus daceloae
Tinaminyssus halcyonus
Tinaminyssus halcyonus
Tinaminyssus halcyonus
Tinaminyssus haleyonus
Sternostoma cooremanti
Ptilonyssus triscutatus
Sternostoma cooremant
Neoboydaia merops
Sternostoma cuculorum —
Sternostoma cuculorum
Sternostoma cuculorum
Ptilonyssus pittae
Ptilonyssus angrensis |
Ptilonyssus echinatus |
Boydaia hirundoae
Ptilonyssus emberizae
Ptilonyssus echinatus
Ptilonyssus langei
Sternostoma tracheacolum
Boydaia hirundoae
Cheramoeca leucosterna (Gould)
Hylochelidon nigricans (Vieillot)
Hylochelidon ariel (Gould)
Microeca fascinans (Latham)
Microeca flavigaster Gould
Petroica rosea Gould
Petroica goodenovit (Vigors and
Horsfield)
Melanodryas cucullata (Latham)
Rhipidura fuliginosa (Sparrman)
Rhipidura setosa
Gaimard)
(Quoy
Rhipidura rufifrons (Latham)
Rhipidura leucophrys (Latham)
Mytagra rubecula (Latham)
Seisura inquieta (Latham)
Arses kaupi Gould
Piezorhynchus alecto (Temminck)
Monarcha melanopsis (Vieillot)
Monarcha trivirgata (Temminck)
Carterornis leucotis (Gould)
Eopsaltria capito Gould
Eopsaltria chrysorrhoa Gould
Pieropodocys maxima (Riippell)
Coracina novaehollandiae (Gmelin)
Coracina hypoleuca (Gould)
Coracina papuensis (Gmelin)
Coracina lineata (Swainson)
Coracina robusta (Latham)
Edoliisoma tenwirostre (Jardine)
Lalage tricolor (Swainson)
Lalage leucomela (Vigors
Horsfield)
Orthonyx temminckii Ranzani
Pomatostomus temporalis (Vigors
and Horsfield)
Smicrornis brevirostris (Gould)
Smicrornis flavescens Gould
Gerygone olivacea (Gould)
Gerygone fusca (Gould)
Gerygone richmondii (Mathews)
Gerygone mouki Mathews
Gerygone palpebrosa Wallace
Cinclorhamphus mathewsi Iredale
Acrocephalus australis (Gould)
Cisticola exilis (Vigors and Hors-
field)
Chthonicola sagittata (Latham)
ROBERT DOMROW
TABLE 4—Continued
White-backed swallow
Tree-martin
Fairy martin
Muscicapidae
Jacky winter
Lemon-breasted flycatcher
Rose-robin
Red-capped robin
‘Hooded robin
Grey fantail
Northern fantail
Rufous fantail
Willie wagtail
Leaden flycatcher
Restless flycatcher
Australian pied flycatcher
Shining flycatcher
Black-faced flycatcher
Spectacled flycatcher
White-eared flycatcher
Pale-yellow robin
Northern yellow robin
Campophagidae
Ground cuckoo-shrike
Black-faced cuckoo-shrike
White - breasted cuckoo -
shrike
Papuan cuckoo-shrike
Barred cuckoo-shrike
Little cuckoo-shrike
Jardine caterpillar-eater
White-winged triller
Varied triller
Timaliidae
Southern chowchilla
Grey-crowned babbler
Sylviidae
Brown weebiil
Yellow weebill
White-throated warbler
Western warbler
Brown warbler
Northern warbler
Black-throated warbler
Rufous songlark
Australian reed-warbler
-Fantail-warbler
Speckled warbler
Ptilonyssus echinatus
Boydaa hirundoae
Ptilonyssus microecae
Ptilonyssus microecae
Ptilonyssus motacillae
Ptilonyssus rhipidurae
Ptilonyssus setosae
Ptilonyssus setosae
Ptilonyssus maccluret
Sternostoma cuculorum
Ruandanyssus terpsiphonet
Ptilonyssus terpsiphonet
Sternostoma cuculorum
Ptilonyssus terpsiphoner
Ptilonyssus terpsvphonetr
Sternostoma cuculorum
Ptilonyssus terpsvphonet
Ptilonyssus terpsiphonet
Ruandanyssus terpsiphoner
Ptilonyssus terpsiphonet
Ptilonyssus monarchae
Sternostoma cuculorum
Ptilonyssus terpsiphonet
Ptilonyssus dioptrornis
Sternostoma cuculorum
Ruandanyssus terpsiphonetr
Ruandanyssus terpsiphonetr
Ruandanyssus terpsiphonet
Ptilonyssus cractict
Boydaia crassipes
Ptilonyssus orthonychus
Passerrhinoptes pomatostomi
Piilonyssus gery ,onae
Ptilonyssus bradypterv
Ptilonyssus acrocephali
Sternostoma tracheacolum
413
414
TABLE 4—Continued
THE NASAL MITES OF QUEENSLAND BIRDS
List of Australian birds examined and their recorded nasal mite parasites—Continued
Acanthiza pusilla (Shaw)
Acanthiza uropygialis Gould
Acanthiza chrysorrhoa (Quoy and
Gaimard)
Acanthiza reguloides Vigors and
Horsfield
Sericornes lathami (Stephens)
Sericornis frontalis (Vigors and
Horsfield)
Sericornis magnirostris (Gould)
Malurus melanotus Gould
Malurus lambertt Vigors and
Horsfield
Malurus amabilis Gould
Malurus melanocephalus (Latham)
Artamus superciliosus (Gould)
Artamus personatus (Gould)
Artamus cyanopterus (Latham)
Artamus cinereus Gould
Ariamus minor Vieillot
Grallina cyanoleuca (Latham)
Colluricincla harmonica (Latham)
Colluricincla megarhyncha (Quoy
and Gaimard)
Pachycephala pectoralis (Latham)
Pachycephala rufiventris (Latham)
Pachycephala griseiceps Gray
Psophodes olivaceus (Latham)
Falcunculus frontatus (Latham)
Oreoica gutturalis (Vigors and
Horsfield)
Turdus merula Linnaeus
Turdus philomelos Brehm
Oreocincla lunulata (Latham)
Epthianura tricolor Gould
Gymnorhina tibicen (Latham)
Cracticus nigrogularis (Gould)
Cracticus torquatus (Latham)
Cracticus quoyi (Lesson
Garnot)
Cracticus mentalis Salvadori and
d’Albertis
Strepera graculina (Shaw)
and
Neositta chrysoptera (Latham)
Neositta leucocephala (Gould)
Neositta striata (Gould)
Brown thornbill
Chestnut-tailed thornbill
Yellow-tailed thornbill
Buff-tailed thornbill
Yellow-throated scrub-wren
White-browed sc: ub-wren
Large-billed scrub-wren
Black-backed blue wren
Variegated wren
Lovely wren
Red-backed wren
Artamidae
White-browed wood-swallow
Masked wood-swallow
Dusky wood-swallow
Black-faced wood-swallow
Little wood-swallow
Grallinidae
Magpie-lark
Pachycephalidae
Grey shrike-thrush
Rufous shrike-thrush
Golden whistler
Rufous whistler
Grey whistler
Faleunculidae
Eastern whipbird
Eastern shrike-tit
Crested bellbird
Turdidae
Blackbird
Song-thrush
Australian ground-thrush
Crimson chat
Cracticidae
Black-backed magpie
Pied butcher-bird
Grey butcher-bird
Black butcher-bird
Black-backed butcher-bird
Pied currawong
Sittidae
Orange-winged sittella
White-headed sittella
Striated sittella
Ruandanyssus terpsiphoner
Boydaia maluri
Ptilonyssus maluri
Boydaia maluri
Ruandanyssus terpsiphoner
Ruandanyssus artami
Ruandanyssus artami
Ptilonyssus grallinae
Ptilonyssus colluricinclae
Ptilonyssus colluricinclae
Sternostoma cuculorum-
Ruandanyssus terpsiphoner
Piilonyssus colluricinclae
Ruandanyssus terpsiphoner
Ptilonyssus motacillae
Ruandanyssus. terpsiphoner
Ptilonyssus colluricinclae
Ptilonyssus psophodae
Sternostoma cuculorum
Sternostoma technaut
Sternostoma technaui
Sternostoma durent
Sternostoma technaur
Ptilonyssus cracticr
Sternostoma thienpontr
Ptilonyssus cractici
Sternostoma thienpontr
Ptilonyssus cractice
Sternostoma thienpontr
Ptilonyssus cracticr
Ptilonyssus cracticr
Sternostoma neosittae
ROBHRT DOMROW
Taste 4—Continued
List of Australian birds examined and their recorded nasal mite parasites—Continued
Climacteris picumnus 'Temminck
Climacteris melanota Gould
Climacteris leucophaea (Latham)
Zosterops lateralis (Latham)
Dicaeum hirundinaceum (Shaw)
Pardalotus ornatus (Gmelin)
Pardalotus punctatus (Shaw)
Pardalotus rubricatus Gould
Pardalotus melanocephalus Gould
Cyrtostomus frenatus (Muller)
Melithreptus albogularis Gould
Plectorhyncha lanceolata Gould
Myzomela sanguinolenta (Latham)
Myzomela pectoralis Gould
Myzomela obscura Gould
Acanthorhynchus tenutrostris
(Latham)
Gliciphila fasciata Gould
Gliciphila indistincta (Vigors and
Horsfield)
Conopophila albogularis (Gould)
Meliphaga fusca Gould
Meliphaga lewinii Swainson
Meliphaga notata (Gould)
Meliphaga gracilis (Gould)
Meliphaga chrysops (Latham)
Meliphaga macleayana (Ramsay)
Meliphaga penicillata Gould
Meliphaga flava (Gould)
Stomiopera unicolor (Gould)
Myzantha melanocephala
(Latham)
Myzantha flavigula (Gould)
Anthochaera chrysoptera (Latham)
Acanthagenys rufogularis Gould
Entomyzon cyanotis (Latham)
Philemon corniculatus (Latham)
Philemon citreogularis (Gould)
Anthus australis Vieillot
Motacilla flava Linnaeus*
Mirafra javanica Horsfield
Certhiidae
Brown tree-creeper
Black tree-creeper
White-throated tree-creeper
_ Zosteropidae
Grey-backed silvereye
Dicaeidae
Mistletoe-bird
Red-tipped pardalote
Spotted pardalote
Red-browed pardalote
Black-headed pardalote
Nectariniidae
Yellow-breasted sunbird
Meliphagidae
White-throated honeyeater
Striped honeyeater
Scarlet honeyeater
Banded honeyeater
Dusky honeyeater
Eastern spinebill
White-breasted honeyeater
Brown honeyeater
Rufous-banded honeyeater
Fuscous honeyeater
Lewin honeyeater
Lesser Lewin honeyeater
Graceful honeyeater
Yellow-faced honeyeater
Macleay honeyeater
White-plumed honeyeater
Yellow honeyeater
White-gaped honeyeater
Noisy miner
Yellow-throated miner
Little wattle-bird
Spiny-cheeked honeyeater
Blue-faced honeyeater
Noisy friar-bird
Little friar-bird
Motacillidae
Australian pipit
Yellow wagtail
Alaudidae
Horsfield bushlark
Ptilonyssus sittae
Ptilonyssus ruandae
Sternostoma cuculorum
Boydaia zosteropis
Ptilonyssus dicaei
Ptilonyssus cinnyris
Sternostoma tracheacolum
Ptilonyssus meliphagae
Ptilonyssus philemoni
Ptilonyssus myzomelae
Ptilonyssus lymozemae
Boydaia myzomelae
Ptilonyssus gliciphilae
Boydaia spatulata
Ptilonyssus
Ptilonyssus
myzomelae
gliciphilae
Sternostoma gliciphilae
Pitilonyssus
Ptilonyssus
Ptilonyssus
Ptilonyssus
Pitilonyssus
Ptilonyssus
Ptilonyssus
Ptilonyssus
Ptilonyssus
Ptilonyssus
Ptilonyssus
Ptilonyssus
Ptilonyssus
Ptilonyssus
Ptilonyssus
Ptilonyssus
Ptilonyssus
Ptilonyssus
Ptilonyssus
thymanzae
philemoni
thymanzae
philemoni
thymanzae
meliphagae
balimoensis
thymanzae
stomioperae
stomioperae
myzanthae
thymanzae
myzanthae
thymanzae
myzanthae
thymanzae
thymanzae
philemoni
philemoni
Boydaia spatulata
Ptilonyssus
philemoni
Boydaia spatulata
Ptilonyssus
motacillae
Ptilonyssus motacillae
Boydaia crassipes
Piilonyssus
capitatus
415
416 THE NASAL MITES OF QUEENSLAND BIRDS
TABLE 4—Continued
Inst of Australian birds examined and their recorded nasal mite parasites—Continued
Chloris chloris (Linnaeus)
Carduelis carduelis (Linnaeus)
Passer montanus (Linnaeus)
Passer domesticus (Linnaeus)
Serinus canaria (Linnaeus)*
Taeniopygia castanotis (Gould)
Steganopleura bichenovit (Vigors
and Horsfield)
Donacola castaneothorax (Gould)
Aidemosyne modesta (Gould)
Aegintha temporalis (Latham)
Bathilda ruficauda (Gould)
Neochmia phaeton (Hombron and
Jacquinot)
Poephila atropygialis Diggles
Poephila personata Gould
Poephila gouldiae (Gould)
Lonchura punctulata (Linnaeus)*
Padda oryzivora (Linnaeus)*
Sturnus vulgaris Linnaeus
Acridotheres tristis (Linnaeus)
Aplonis metallica (Temminck)
Oriolus sagittatus (Latham)
Oriolus flavocinctus (King)
Sphecotheres vieillotia Vigors and
Horsfield
Sphecotheres flaviventris Gould
Chibia bracteata (Gould)
Ptiloris paradiseus Swainson
Pitiloris victoriae Gould
Ptiloris magnificus (Vieillot)
Ptilonorhynchus violaceus
(Vieillot)
Ailuroedus crassirosiris (Paykull)
Ailuroedus melanotus (Gray)
Chlamydera maculata (Gould)
Chlamydera mnuchalis (Jardine
and Selby)
Sericulus chrysocephalus (Lewin)
Corvus coronoides Vigors and
Horsfield
Corvus cecilae Mathews
Struthidea cinerea Gould
Corcorax melanorhamphus
(Vieillot)
Fringillidae
Greenfinch
Goldfinch Q
Tree-sparrow
House-sparrow
Canary
Ploceidae
Zebra finch
Banded finch
Chestnut-breasted finch
Plum-headed finch
Red-browed firetail
Star finch
Crimson finch
Black-tailed finch
Masked finch
Gouldian finch
Spice finch
Java sparrow
Sturnidae
Starling
Common myna
Australian shining starling
: Oriolidae
Olive-backed oriole
Yellow oriole
Southern figbird
Yellow figbird
Dicruridae
Spangled drongo
Paradiseidae
Paradise rifle-bird
Victoria rifle-bird
Magnificent rifle-bird
Ptilonorhynchidae
Satin bower-bird
Green catbird
Spotted catbird
Spotted bower-bird
Great bower-bird
Regent bower-bird
Corvidae
Australian raven
Australian crow
Apostle-bird
White-winged chough
Ptilonyssus carduelis
Ptilonyssus pygmaeus
Ptilonyssus motacillae
Sternostoma tracheacolum
Boydaia crassipes
Ptilonyssus nudus
Ptilonyssus hirsti
Ptilonyssus nudus
Sternostoma tracheacolum
Sternostoma cryptorhynchum
Boydaia crassipes
Sternostoma tracheacolum
Ptilonyssus motacillae
Ptilonyssus neochmiae
Ptilonyssus emberizae
Sternostoma tracheacolum
Ptilonyssus motacillae
Sternostoma paddae
Ptilonyssus elbeli
Boydaia sturni
Ptilonyssus elbeli
Ptilonyssus motacillae
Boydaia sturni
Ptilonyssus sturnopastoris
Pitilonyssus trowessarte
Ptilonyssus trouessarti
Ptilonyssus sphecotheris
Ptilonyssus sphecotheris
Ptilonyssus dicrurt
Sternostoma thienponti
Ptilonyssus novaeguineae
Pitilonyssus arluroedi
Ptilonyssus sphecotheris
Ruandanyssus terpsiphoner
Piilonyssus struthideae
Ptilonyssus corcoracis
ROBERT DOMROW 417
Acknowledgements
The birds examined during this study were collected under permit from
the Queensland Department of Primary Industries. The Queensland Museum
kindly identified the species not well known to me, and Mr. H. J. Larvey
(Animal Health Station, Oonoonba) furnished data from his unpublished
check-list of Queensland birds. Of the many collectors listed in Section I
above, my assistants, Messrs. I. D. Fanning, B. H. Kay, and J. S. Welch.
deserve special mention for their companionship in the field and assistance
in the laboratory. Dr. K. H. L. Key (Division of Entomology, C. 8. T. R. O.,
Canberra) offered advice on nomenclatural problems, while Drs. A. Fain,
R. W. Strandtmann, and N. Wilson kindly criticized my manuscript and
placed many specimens at my disposal. Drs. P. L. G. Benoit (Royal Museum
of Central Africa, Tervuren), N. G. Bregetova (Zoological Institute, U.S. 8. R.
Academy of Sciences, Leningrad), and F. Zumpt (South African Institute
of Medical Research, Johannesburg) also kindly loaned or donated specimens.
Publication costs were partially met from W. H. O. support to the Arboyirus
Reference Laboratory functions of this Institute. Miss D. Lees patiently
prepared the typescript, and Mr. F. B. Carter the photographs, while
Mrs. M. Macgregor provided me with copies of all the early references.
I am most grateful to them all.
References
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418 THE NASAL MITES OF QUEENSLAND BIRDS
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ROBERT DOMROW 419
, 1957e.—Les acariens des familles Epidermoptidae et Rhinonyssidae parasites
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a new genus and five new species. J. ent. Soc. sth Afr., 22: 18-34.
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sud-americanis (Rhinonyssidae: Mesostigmata). Emendation du nom Neoboydaia
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33: 3-12.
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(Acarina : Mesostigmata et Trombidiformes). Revue Zool. Bot. afr., 70: 29-39.
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d’oiseaux (Mesostigmata: Rhinonyssidae). Revue Zool. Bot. afr., 70: 123-128.
, 1964e—Les Ereynetidae de la collection Berlese a Florence. Designation
d’une espéce type pour le genre Hreynetes Berlese. Redia, 49: 87-111
, 1965a.—Quelques aspects de l’endoparasitisme par les acariens. Annis Parasit.
hum. comp., 40: 317-327.
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72: 152-160.
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420 THE NASAL MITES OF QUEENSLAND BIRDS
, 1966b.—Notes sur quelques Rhinonyssidae (Acari: Mesostigmata). Revue
Zool. Bot. afr., 74: 83-96.
, 1967a.—Un acarien remarquable récolté sur un tarsier (Heterocoptidae
f. n.: Sarcoptiformes). Zool. Anz., 178: 90-94.
, 19676.—Trois nouveaux Rhinonyssidae avec note sur la nymphiparite
dans cette famille. Revue Zool. Bot. afrs 76: 149-156.
Fain, A., and AITKEN, T. H. G., 1967.—Les acariens parasites nasicoles des oiseaux
de Trinidad (Indes Occidentales). I. Rhinonyssidae: Mesostigmates. Bull. Inst.
r. Sci. nat. Belg., 43: 1-44.
Fain, A., and BArort, J., 1963.—Conjonctivite epizootique chez Padda oryzivora produite
par un acarien Sternostoma paddae Fain, 1958. Bull. Soc. r. Zool. Anvers, 31: 3-6.
Fain, A., and CARpPENTIER, J., 1958—Acariase pulmonaire mortelle chez des canaris
du Zoo. Bull. Soc. r. Zool. Anvers, 9: 21-24.
Fain, A., and Hyranp, K. E., 1962a—The mites parasitic in the lungs of birds. The
variability of Sternostoma tracheacolum Lawrence, 1948, in domestic and wild birds.
Parasitology, 52: 401-424.
—_———_, ————_., 1962b.— On three species of rhinonyssids described by Hirst, Ann.
Mag. nat. Hist., (13) 5 : 341-348.
Fain, A., and JouHnston, D. E., 1966.——Nouveaux acariens nasicoles d’oiseaux nord-
americains (Acari: Rhinonyssidae). Bull. Soc. r. Zool. Anvers, 38: 25-41.
Farin, A., and Morrermans, J., 1959—Sur la présence d’un nouvel halarachnide chez
un manchot papou (Acarina: Mesostigmata). Bull. Soc. r. Zool. Anvers, 12: 21-27.
Fain, A. and NAapcHATrRAM, M., 1962.—Acariens nasicoles de Malaisie. I. Hreynetoides
malayi n. g., nN. sp., parasite d’un nectarin (Hreynetidae : Trombidiformes). Z.
ParasitKde, 22: 68-82.
Forp, H. G., 1959.—Boydaia tyrannis n. sp. (Acarina, Speleognathidae), a new mite
from the nasal cavity of the eastern kingbird, Tyrannus tyrannus (Linnaeus).
Trans. Am. microse. Soc., 78: 379-385.
Furman, D. P., 1957.--Revision of the genus’ Sternostoma Berlese and Trouessart
(Acarina : Rhinonyssidae). Hilgardia, 26: 473-495.
, 1963.—Problems in the control of poultry mites. Adv. Acar., 1: 30-38.
Georce, J. E., 1961—The nasal mites of the genus Ptilonyssus (Acarina: Rhinonys-
sidae) occurring in some North American passeriform birds. J. Kans. ent. Soc.,
34: 105-132.
GIEBEL, C., 1871.—Ueber einige Milben. Z. ges. Naturw., 38: 29-32.
GRETILLAT, S., CApron, A., and Bryeoo, E. R., 1959—Acariens Rhinonyssidae de Mada-
gascar. Agapornyssinae, n. sfam.; Agapornyssus, n. g.; Agapornyssus faini Nn. Sp.;
Ptilonyssus madagascariensis n. sp. et Neonyssus marcandrei n. sp., parasites des
fosses nasales et des poumons d’oiseaux malgaches. Acarologia, 1: 375-384.
Hirst, S., 1921a¢—On some new or little-known Acari, mostly parasitic in habit. Proc.
zool. Soc. Lond., 1921: 357-378.
, 19216.—On some new parasitic mites. Proc. zool. Soc. Lond., 1921: 769-802.
, 1923.—On some new or little-known species of Acari. Proc. zool. Soc. Lond.,
1923 : 971-1000.
HYLAnp, K. E., 1961—~Sternostoma longisetosa [sic], a new species of nasal mite from
the eastern kingbird with notes on the occurrence of Tyranninyssus spinosus Brooks
and Strandtmann in southern Michigan (Acarina; Rhinonyssidae). Acarologia,
3: 279-284.
, 1963.—Current trends in the systematics of acarines endoparasitic in verte-
brates. Adv. Acar., 1: 365-373.
Imus, A. D., 1957—‘“A general Textbook of Entomology including the Anatomy,
Physiology, Development and Classification of Insects”. 9th Ed., revised by
O. W. Richards and R. G. Davies. (Methuen: London.)
Kerast, A., 1957—Variation and speciation in the genus Climacteris Temminck (Aves :
Sittidae). Aust. J. Zool., 5: 474-495.
, 1961.—Bird speciation on the Australian continent. Bull. Mus. comp. Zool.
Harv., 123: 305-495.
KirnKwoop, A., 1963.—Longevity of the mites Dermanyssus gallinae and Liponyssus
Ssylviarum. Expl Parasit., 14: 358-366.
LAWRENCE, R. F., 1956.—Studies on South African fur-mites (Trombidiformes and
Sarcoptiformes). Ann. Natal Mus., 13: 337-375.
Leacu, J. A., 1958—‘‘An Australian Bird Book. A complete Guide to the Birds of
Australia”. 9th Hd., revised by P. C. Morrison (Whitcombe and Tombs: Melbourne
etc.)
, et al., 1926—“‘Official Checklist of the Birds of Australia”. 2nd Ed. (Royal
Australian Ornithologists’ Union : Melbourne.)
Linpquist, EH. E., and Evans, G. O., 1965.—Taxonomic concepts in the Ascidae, with a
modified setal nomenclature for the idiosoma of the Gamasina /{Acarina:
Mesostigmata). Mem. ent. Soc. Canada, 47: 1-64.
ROBERT DOMROW 421
LompBarpini, G., 19386.—Elenco alfabetico di specie esistenti nell’acaroteca della R
Stazione di Entomologia Agraria di Firenze. Redia, 22: 37-51.
Maa, T. C., and Kuo, J. S., 1965.—A field survey of arthropod parasites of birds in
Taiwan. J. med. Ent., 1: 395-401.
McCuure, H. E., 1963.—English vernacular names of the birds of the Malaysian subregion
Malay. Nat. J., 17: 76-121.
Maruery, W. J., 1967.—Respiratory acariasis due to Sternostoma tracheacolum in the
budgerigar. J. Am. vet. med. Ass., 150: 777-780.
Mayr, E., and Amapon, D., 1951.—A classification of recent birds. Am. Mus. Novit.,
1496: 1-42.
Mayr, E., Linstry, E. G., and Usinerr, R. L., 1953.—‘Methods and Principles of
systematic Zoology”. (McGraw-Hill: New York etc.)
Mircnetyt, R. W., 1963.—Comparative morphology of the life stages of the nasal mite
Sternostoma rhinolethrum (Mesostigmata: Rhinonyssidae). Aust. vet. J., 42 : 262-264
Morray, M. D., 1966.—Control of respriatory acariasis of Gouldian finches caused by
Sternostoma rhinolethrum (Mesostigmata : Rhinonyssidae). Aust. vet. J., 42:262—-264.
NeAvr, S. A., 1939-40.—‘‘Nomenclator zoologicus. A List of the Names of Genera and
Subgenera in Zoology from the tenth Edition of Linnaeus 1758 to the End of 1935”.
(Zoological Society of London.)
PARKER, T. J., and HAasweiit, W. A., 1962.—“Textbook of Zoology’. Vol. II, 7th Ed.,
revised by A. J. Marshall. (Macmillan : London.)
Pereira, C., and pr CaAsrro, M. P., 1949.—Revisao da subfamilia “Ptilonyssinae Castro,
1948” (Acari Mesostigmata : Rhinonyssidae Vitz.), com a descricao de algumas
espécies novas. Archos Inst. biol., S Paulo, 19: 217-235.
SAKAKIBARA, I., 1967.—New nasal mites, Ptilonyssus and Paraneonyssus (Acarina :
Mesostigmata), from Taiwan and New Guinea. Pacif. Insects, 9: 597-601.
Setinick, M., 1965—Epistom und Tectum bei den Mesostigmata (Acarina). Acarologia,
7: 594-597.
, 1967.—“Onece more Tectum and Epistome’’. (Privately printed: Grosshans-
dorf.) :
Sranprast, H. A., 1965.—Notes on the birds of Mitchell River. Qd Nat., 17: 91-94.
Stott, N. R., et al., 1964—‘‘International Code of zoological Nomenclature adopted
by the XV International Congress of Zoology”. 2nd Ed. (International Trust for
Zoological Nomenclature : London.)
STRANDTMANN, R. W., 1951.—The mesostigmatic nasal mites of birds. II. New and
poorly known species of Rhinonyssidae. J. Parasit., 37: 129-140.
, 1956.—The mesostigmatic nasal mites of birds. IV. The species and hosts
of the genus Rhinonyssus (Acarina, Rhinonyssidae). Proc. ent. Soc. Wash.,
58: 129-142.
————, 1958.—Host specificity of bird nasal mites (Rhinonyssidae) is a function
of the gregariousness of the host. Proc. X int. Congr. Ent., 1: 909-911.
, 1959.—New records for Rhinonyssus himantopus and notes on other species
of the genus. J. Kans. ent. Soc., 32: 133-136.
, 1961.—The immature stages of the Ptilonyssus complex (Acari : Mesostigmata:
Rhinonyssidae). Proc. XI int. Congr. Ent., 1: 283-286.
, 1962.—A ptilonyssid mite from the sparrow hawk, Falco sparverius (Acarina:
Rhinonyssidae). Proc.- ent. Soc. Wash., 64: 100-102.
STRANDTMANN, R. W., and FurmMAn, D. P., 1956.—A new species of mite, Paraneonyssus
icteridius, from the nasal cavities of blackbirds. Pan-Pacif. Ent., 32: 167-173.
STRANDTMANN, R. W., and WHARTON, G. W., 1958—A manual of mesostigmatid mites
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Uppsala.). Reprinted 1905 as Vol. IV of “Fauna arctica’. (Gustav Fischer: Jena.)
TROUESSART, E., 1893.—Note sur les sarcoptides pilicoles (Listrophorinae). C. r. Séane.
Soc. Biol., (9) 5: 698-700.
— , 1894-—Note sur les acariens parasites des fosses nasales des oiseaux. C. 1.
Séanc. Soc. Biol., (10) 1: 723-724.
, 1895.—Note sur un acarien parasite des fosses nasales de l’oie domestique
(Sternostomum rhinolethrum, n. sp.). Revue Sci. nat. appl., 42 : 392-394.
, 1896.—Description d’un genre nouveau (Labidocarpus) et de deux espéces
nouvelles de sarcoptides pilicoles. Bull. Soc. ent. Fr., 1895: Ixxxii-lxxxvii.
TROUESSART, E., and NEuMANN, G., 1890.—Un type nouveau de sarcoptides plumicoles,
le Chirodiscus amplexans, g. n., sp. n. Bull. scient. Fr. Belg., 22: 392-398.
VitztHum, H., 1935.—Milben aus der Nasenhdhle von Voégeln. J. Orn., Lpz., 83: 563-587.
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(Becker and Erler: Leipzig.)
Wess, L. J., 1959-——A physiognomic classification of Australian rain forests. J. Ecol.,
47: 551-570.
422 THE NASAL MITES OF QUEENSLAND BIRDS
WHITEHEAD, R. H., Douerty, R. L., Domrow, R., Stanprast, H., and WertTers, E. J.,
1968.—Studies of the epidemiology of arthropod-borne virus infections at Mitchell
River Mission, Cape York Peninsula, North Queensland. III. Virus studies of
wild birds, 1964-1967. Trans. r. Soc. trop. Med. Hyg., 62: 489-445.
Witson, N., 1964—New records and descriptions of Rhinonyssidae, mostly from New
Guinea (Acarina: Mesostigmata). Pacif. Insects, 6: 357-388.
, 1965—New records and descriptions of Reallinyssus from Pacific birds
(Acarina: Mesostigmata). Pacif. Insects, 7: 623-639.
, 1966a.—Mesonyssus (Acarina: Mesostigmata) from New Guinea, Philippine
and Taiwan birds. Pacif. Insects, 8: 601-609.
, 1966b.—New records and a new species of Mesonyssus (Acarina: Mesostig-
mata) from southeast Asian Psittacidae (Aves: Psittaciformes). Pacif. Insects,
8: 759-769.
, 1967.—Rallinyssus from Philippine birds (Acarina: Mesostigmata). Philipp.
J. Sci., 95: 215-226.
, 1968a.—New records and a new species of Mesonyssus (Mesostigmata :
Rhinonyssidae) from parrots (Psittaciformes, Psittacidae). J. Parasit., 54: 395-401.
, 1968b.—Records of nasal mites (Mesostigmata: Rhinonyssidae) from New
Guinea, Philippines and United States. J. med. Ent., 5: 211-223.
Womerstey, H., 1936.—On a new family of Acarina, with description of a new genus
and species. Ann. Mag. nat. Hist., (10) 18: 312-315.
, 1948—A modification of Berlese’s medium for the microscopic mounting of
Acarina and other small arthropods. Trans. R. Soc. S. Aust., 67: 181-182.
, 1953—A new genus and species of Speleognathidae (Acarina) from South
Australia. Trans. R. Soc. 8S. Aust., 76: 82-84.
Woottey, T. A., 1961.—A review of the phylogeny of mites. A. Rev. Hnt., 6: 263-284.
Youne, J. Z., 1950—‘“The Life of Vertebrates’. (Clarendon Press: Oxford.)
ZumMpt, F., and Patterson, P. M., 1951.—Further notes on laelaptid mites parasitic on
vertebrates. A preliminary study to the Ethiopian fauna. J. ent. Soe. sth. Afr.,
14: 63-93.
Zoumpt, F., and Titt, W. M., 1955—Nasal mites of birds hitherto known from the
Ethiopian region, with keys and descriptions of nine new species (Acarina:
Laelaptidae). J. ent. Soc. sth. Afr., 18 : 60-92.
; , 1961.—Suborder Mesostigmata. Publs S. Afr. Inst. med. Res., 9: 17-91.
ADDENDA
Recent visits to Kowanyama have necessitated the following changes to
the list of birds recorded there by Standfast (1965) and Domrow (1967a).
Delete Dupetor flavicollis and Ptilonorhynchus violaceus as misidentifications
of Nycticorax caledonicus (juvenile) and Hudynamys orientalis, respectively.
Add Tringa nebularia (D), Gallinago hardwickii (W), Hamirostra melanos-
terna (D), Acrocephalus australis (W),,and Anthus australis (D). Alectura
lathami and Porphyrio melanotus are confirmed as occurring in the area,
where 139 species of birds are now known.
Two recent papers by Sakakibara should be noted (1968, J. med. Ent.,
5 : 298-309 ; 1968, Acarologia, 10: 426-431). His description of the exact nature
of the dorsal shields of Ptilonyssus novaeguineae (Hirst), which is keyed out
above on the assumption that it possesses only the podonotal and pygidial
shields typical of the vast majority of species, allows its more exact placement
in couplet 12, which includes P. triscutatus (Vitzthum) and P. sittae Fain.
It is immediately separable by the shape and proportions of its three dorsal
shields and the nature of the leg setation.
I would place P. paradisaeus Sakakibara as a synonym of P. novae-
guineae—the minor setational differences alleged are essentially in characters
prone to considerable variation in this group, and both are from paradiseids
(Sakakibara lists grackles, Icteridae, among the hosts, but this neotropical
family does not occur in New Guinea—this recalls Womersley and Audy’s
ROBERT DOMROW 423
caution (1957, Stud. Inst. med. Kes. I'.M.S., 28: 231-296) on the tendency
for the laconic “rat” and “mouse” on field labels later to be interpreted as
“Rattus” and “Mus”.
Likewise, I would place P. trouessarti pseudotrouessarti Sakakibara as
a synonym of P. trouessarti (Hirst). Sakakibara also recognizes P. orioli
Fain, but his illustration of a paratype is at variance with Fain’s, and
until the holotypes are compared, I maintain P. orioli under the synonymy
of P. trouessarti. Oriolids figure largely among the hosts of these taxa.
Two other species discussed above (P. terpsiphonei Fain and P. estril-
dicola Fain) are also recorded from New Guinea.
P. missimensis Sakakibara is the form of P. philemoni Domrow described
and figured above from Meliphaga spp., and is accordingly synonymized
under that species.—March 5, 1969.
Messrs. E. T. Bulfin, A. L. Dyce, H. A. S., and R. D. collected these
novelties (not entered in Tables) at Kowanyama in April 1969:
Rhinonyssus comventris, 4292, 16, Hrolia ruficollis (the variety
described with ¢ differing from @ only in primary sexual characters).
Rhinoecius cooremani, 32 2, Ninox connivens (deep in nares) .*
Ptilonyssus neochmiae, 12, Neochmia albiventer Mathews.* +i
Ptilonyssus echinatus, 12, Hylochelidon ariel (pygidial shields small,
well separated) .*+
Neoboydaia psittaculae, 12, Aprosmictus erythropterus (differs from
Fain’s chaetotaxy in showing femora 4.3.2.3; specimen from T'richoglossus
shows femora 4/3.2.2.2 and tibiae 5.3.2.3; specimen from Psitteuteles shows
tibiae 4.2.2.2) .*
G. et sp. indet. (Speleognathinae), 1 29, Turnig velox (Gould) .*7i
Mycteroptes basilewskyi, 42 9, Hurystomus orientalis (* represents
new host record, + bird new to Table 4, and ¢ bird new to Kowanyama).
Additional species clearly sighted by R. D. at Kowanyama are Numenius
phaeopus, Erolia ferruginea, Ardea sumatrana Raffles, Dupetor flavicollis
(Latham), Tyto alba, and Artamus minor, making a total of 148.
Finally, Sternostoma tracheacolum caused respiratory symptoms in
aviary-bred Serinus canaria, Burnie, Tas., iv. 1969, R. W. Mason and A. Little.
Sternostoma borceanum Feider and Mironescu, 1968, Anal. stiint. Univ.
Al. I, Cuza, 14: 105, from various European turdids, is an obvious synonym
of S. technau.—May 27, 1969.
424 THE NASAL MITES OF QUEENSLAND BIRDS
INDEX OF FAMILIES, GENERA, AND Species or Mires DIscusseD
(Accepted names in italics; synonyms in roman, with original termination)
Page
accipitris (Ophthalmognathus) 390
acrocephali Br eae .. 308
Agapornyssus.. .. O04
ailuroedi (Ptilonyssus) .. 348
alcippei (Ptilonyssus) . .. 366
amaurornis ( Rallinyssus ) .. 323
angrensis (Ptilonyssus) .. 048
aprosmicti (Tinaminyssus) .. 306
ardeae (Tinaminyssus) oreatestl al
artami (Ruandanyssus) .. 330
Astridiella .. 335
aureliant (Neoboydaia) .- 389
Aureliania .. 388
balimoensis (Ptilonyssus) .. 368
basilewskyi (Mycteroptes) .. 395
batis (Sternostoma) .. ot
belopolsku (Tinaminyssus) .. 311
benoiti (Larinyssus) .. ae ay
benoit ee da ee ba BIS
Boydaia a 380, 401
boydi (Ster nostoma) oc cp ote)
bradypteri (Ptilonyssus) . .. 358
bruxellarum (Ptilonyssus) .. 348
bubulci (Tinaminyssus) sii
capitatus (Ptilonyssus) Hey oss,
caprimulgi (Ptilonyssus) PETS4L9
carduelis (Ptilonyssus) 3 ODS
Cas { .. 335
castroae (Sternostoma) ae OL
caudistigmus (Rallinyssus) .. 324
cerchneis (Ptilonyssus) O74
chalcopsittae (Tinaminyssus) 311
chalybeaedomesticae (Ptilonys-
sus) a 349
charadricola (Speleognathop.
sis) 393
chloris (Ptilonyssus) 2h 55 aa)
cinnyricinch (Ptilonyssus) .. 352
cinnyris (Ptilonyssus) .. 366
colluricinclae (Ptilonyssus) .. 339
columbae (Tinaminyssus ) .. 313
colymbiformi (Neoboydaia) .. 389
congolensis (Rallinyssus) oS
coniwentris (Rhinonyssus ) er orl
cooremani (Rhinoecius) 5) OBS
Page
cooremani (Sternostoma) .. 316
corcoracis (Ptilonyssus) .. 845
cractici (Ptilonyssus) .. .. 349
crassipes (Boydaia) .. . 386
cryptorhynchum (Sternostoma) 379
cuculorum (Sternostoma) .. 376
daceloae (Tinaminyssus) .. d22
dartevellei (Rhinonyssus) .. 325
Dermanyssidae .. st .. 3038
dicaei (Ptilonyssus) .. .. 366
dicruri (Ptilonyssus) .. .. d48
dioptrornis (Ptilonyssus) .. B47
dogieli (Ophthalmognathus) .. 392
domicellae (Tinaminyssus) .. 311
dureni (Sternostoma) .. .. d16
echinatus (Ptilonyssus) Lope BLO
echinipes (Rhinonyssus) segs 216
echongi (Ruandanyssus) ae
elbeli (Ptilonyssus) .. .. 348
emberizae (Ptilonyssus) .. 309
enicuri (Ptilonyssus) .. .. d47
Epidermoptidae .. ; .. 393
epileus (Tinaminyssus) Fale
Ereynetidae 3 .. 380
estrildicola (Ptilonyssus) .. do2
faini (Ptilonyssus) te .. 348
faini (Sternostoma) .. .. OT
. falconis (Boydaia) Ly Sol
Falconyssus ¥ iY .. 305
Flavionyssus y a .. 3835
Frigilonyssus a ase)
fringillicola (Ptilonyssus) .. 302
fulicae (Sternostoma) . ae aiel!)
gall (Speleognathopsis) 4
gallinae (Rhinoptes) .. .. 395
gallinulae (Rallinyssus) .. 020
geopeliae (Tinaminyssus) Piolo
gerygonae (Ptilonyssus) .. 363
gliciphilae (Ptilonyssus) .. 310
gluciphilae (Sternostoma) =A
grallinae (Ptilonyssus) .. 366
haleyoni (Sternostoma) MID
halcyonus (Tinaminyssus ) oa
Hapalognatha .. pee asso 3)
himantopus (Rhinonyssus) 5 a26
ROBERT DOMROW 425
Page Page
hirsti (Ptilonyssus) —.. .. d4l orioli (Ptilonyssus) 373
hirsutus (‘Tinaminyssus ) .. ol2 orthonychus (Ptilonyssus) 354
hirtus (Tinaminyssus) .. SLs Otocorinyssus v; MCB Sh
hirundoae (Boydaia) .. .. d8l Oxleya ys .. 394, 401
ixobrychi (Tinaminyssus ) aera bl paddae (Sternostoma) .. -» OW
kakatuae (Tinaminyssus ) .. 306 Paraneonyssus .. oe .. 800
langei (Ptilonyssus) —.. .. ond Passeronyssus : ve eao
laniorum (Sternostoma ) ALM sarte, Passerrhinoptes .. 094, 401
Larinyssus ae .. O22) OOF Periglischrodes .. Oi: By pp
levinseni (Rhinonyssus ) selec F 3) petiti (Larinyssus) 322
lobatus (Ptilonyssus) .. .. oo2 phabus (Tinaminyssus) ew LT
Locustellonyssus : .. doo philemoni (Ptilonyssus) ASL
lusciniae (P tilonyssus) .. d47 philomachi (Neoboydaia) 388
lymozemae (Ptilonyssus ) oy phoenicuri (Ptilonyssus ) Deoee
macelurei (Ptilonyssus) .. 366 Pipronyssus ot oe .. oa6
macropygiae (Tinaminyssus 319 pittae (Ptilonyssus) .. .. B49
maluri (Boydaia) - .. 386 platycerci (Tinaminyssus) .. 306
maluri (Ptilonyssus) .. 36) Bay) platytricha (Ptilonyssus) ey |
marcandrei (Tinaminyssus) ios) i | pulvialis (Rhinonyssus) .. 226
meddai (Sternostoma) .. hs ened podargi (Oxleya) : .. 394
megaloprepiae (Tinaminyssus) 313 poliocephah (Rhinonyssus) 325
meliphagae (Ptilonyssus) Bee 04 pomatostomi (Passerrhinoptes) 394
melloi (Tinaminyssus) .. ss oli porphyrionis ed we
merops (Neoboydaia) .. Sguatel sis) ; 393
Mesonyssoides (bis) .. .. 305 porzanae (Rallinyssus) . 323
Mesonyssus ot fg .. 305 prima (Ptilonyssus) .. 5 ay
Metaboydaia ie -. 3d09 psittaculae (Neoboydaia) -. 388
microecae (Ptilonyssus) .. 356 Psittanyssus a OUD
minutus (Rhinonyssus ) .. 326 psophodae (Ptilonyssus) .. 366
monarchae (Ptilonyssus) 22 300) pternistis (Rhinoptes) . 2» BEE)
motacillae (Ptilonyssus) -- 352 ptilinopi ( Tinaminyssus) pei,
Mycteroptes oe .. 395, 402 Ptilonyssoides .. ee im. od
myristiciworae (Tinaminyssus) 316 Ptilonyssus : 305, 397
myzanthae (Ptilonyssus) J A100 pygmaeus (Ptilonyssus) .. 341
myzomelae (Boydaia) .. .. 383 Ralliboydaia a : .. 388
myzomelae (Ptilonyssus) . eon) Rallinyssoides.. a 52 320
neglectus (Rhinonyssus) ay Rallinyssus oe ozone (
Neoboydaia se 388, 401 rallus (Rallinyssus) .. .. 323
neochmiae (Ptilony ssus) .. d41 Rhinacarus os Je 334
Neonyssoides ae “is .. 333 Rhinoecius : $3 333, 397
Neonyssus 25 SRS rhinolethrum (Rhinonyssus) 324.
neopsittaci (Tinaminyssus) big) Bulal Rhinonyssinae.. .. 3038, 402
neosittae (Sternostoma) Be Rhinonyssoides .. 2S .. dao
Neospeleognathus ae 5 Gator) Rhinonyssus oie 4S ee BE oe
Neotyranninyssus e: sataaO Rhinoptes ae =. 0895, 402
nigra (Boydaia) ss .. 386 Rhinosterna as Sot
niteschi (Ptilonyssus) .. e. O48 rhipidurae (Ptilonyssus) .- 366
novaeguineae (Ptilonyssus) .. 374 Rochanyssus ce Sao
nudus (Ptilonyssus) .. 372 ruandae (Ptilonyssus) . .. 366
numidae (Schoutedenocoptes) 395 Ruandanyssus.. R. Veco son
nycticoracis (Tinaminyssus) .. 311 schelii (Rhinonyssus) .. . 326
ocyphabus (Tinaminyssus) .. 312 schoutedeni (Ophthalmogna-
Opthalmognathus oR rates sme kD IE thus) .. se OOe
orbicularis (Larinyssus) Ss 322 Schoutedenocoptes a= 89d; 402
J
426 THE NASAL MITES
Page
setosae (Ptilonyssus) .. 362
sittae (Ptilonyssus) .. d48
Sommatericola .. . 324
spatulata (Boydaia) 22 Biel
spatulatum (Sternostoma ) . 380
Speleognathinae .. 380
Speleognathopsis : 392, 401
sphecotheris (Ptilonyssus) . dt2
sphenisci EEO NS, . 326
Spizonyssus .. 30d
Sternoecius .. df4
Sternostoma 374, 400
Sternostomoides : xen om
stomioperae (Ptilonyssus) eroxe!
strandtmanni (Rhinonyssus) .. 326
strandtmanni (Turbinoptes) .. 394
streptopeliae Tinaminyssus) .. 312
striatus (Ophthalmognathus) .. 389
struthideae (Ptilonyssus) . 348
sturmi (Boydaa) ' aerate!)
sturnopastoris (Ptilonyssus) .. 372
taeniopygiae (Ptilonyssus) .. 352
taperaefuscae (Ptilonyssus) .. 349
technaui (Sternostoma) . o19
terpsiphoner (Ptilonyssus ) . 356
OF QUEENSLAND BIRDS
Page
terpsiphonei (Ruandanyssus) .. 329
thienponti (Sternostoma) . 315
thymanzae (Ptilonyssus) youd
Tinaminyssus .. .. 305, 396
tracheacolum (Sternostoma) .. 377
Travanyssus . 305
trichoglossi (Tinaminyssus) . oll
tringae (Rhinonyssus) . mold
triscutatus ( Ptilonyssus) . d47
Trispeleognathus .. 3889
Trochilonyssus . 386
trouessarti (Pritonyssus) . 313
Turbinoptes , 394, 401
Turbinoptinae .. 393
turdi (Sternostoma ) ion
Tyranninyssus .. 335
tytonis (Rhinoecius) .. 333
urolestis (Sternostoma ) . 376
Vitznyssus : tured
wai (Speleootanitaneay . 393
welchi (Tinaminyssus) .. yy elG
zosteropis (Boydma) = OL
zosteropus (Sternostoma) . 376
Zumptnyssus . 338
Proc. Linn. Soc. N.S.W., Vol. 98, Part 3 PLATR XXX
‘bit
ee
Reed .
Plate xxx
Fig. 1. Rain-forest (3,000 ft.) with Wilson’s Peak in background.
Fig. 2. Tropical woodland along Cressbrook Ck., near Esk.
Proc. Linn. Soc. N.S.W., Vol. 98, Part 3 PLATE XXXI
fa) OT Clay i
Plate xxxI
Fig. 1. Semi-arid vegetation and sand dune near Windorah.
Fig. 2. Protected beach flats at low tide, Tin Can Bay.
TYPE SPECIMENS IN THE MACLEAY MUSEUM, UNIVERSITY
OF SYDNEY
II, AMPHIBIANS AND REPTILES
JuDY GoLtpMAN, lL. Hitt and. P. J. STANBURY
School of Biological Sciences, University of Sydney
{Read 30th October, 1968]
Synopsis
The existing amphibian and reptilian type specimens in the Macleay Museum
are listed by families in current phylogenetic order.
INTRODUCTION
The Macleay Museum at the University of Sydney contains a
comprehensive zoological collection including a number of Australian type
specimens. Lists of insect types (Hahn, 1962) and fish types (Stanbury,
1968) have been published. Further lists are being compiled. This paper
lists the existing amphibian and reptilian type specimens in the Macleay
Museum and their current taxonomic status. In all 7 amphibian and 58
reptilian types are catalogued.
The majority of the amphibian and reptilian types were collected on
the “Chevert” expedition led by William Macleay to coastal north Queensland,
Torres Straits and the southern coast of New Guinea in 1875 (Macmillan,
1957). All these were named and described by Macleay (1877a, b, c, d). Other
type material was collected for Macleay by collectors in Australia, but since
his death few types have been lodged in the Museum.
Several Macleay types appear to have been lost. The Museum has had
a varied history (Anderson, 1965) with changes in location during which
some specimens were presumably misplaced. In addition there have been
periods without adequate curatorship when some types, particularly the
geckos, dried out. As a. result, some of the species described by Macleay
have had to be ignored in subsequent taxonomic studies, particularly as
his original descriptions now seem inadequate to define the species.
The types are stored in glass jars in 70% alcohol. The Macleay Museum
Register numbers (R numbers) were entered in 1965 by the previous curator,
Mrs. J. Anderson. These numbers were combined with the numbers given
to some, but not all, specimens by S. J. Copland in 1945. Copland numbers
have been given the prefix MR and where applicable they appear after
the present R numbers in the list of type specimens.
AMPHIBIAN TYPES
From the collection of the “Chevert” expedition, Macleay (1877d)
described five new species of frogs, of which two are lodged in the Macleay
Museum, two in the Queensland Museum, and one, Litoria dorsalis, appears
to have been lost. Neither Boulenger (1882) nor Fry (1913) were able to
classify this species on the basis of Macleay’s description. Apart from
Macleay’s specimens, five other amphibian types are lodged in the Museum.
PROCEEDINGS OF THE LINNEAN Society oF NEw SouTH WALES. VoL. 93, Part 3
428 SPECIMENS IN THE MACLEAY MUSEUM, II. AMPHIBIANS AND REPTILES
REPTILIAN TYPES
ORDER SQUAMATA
Fam. Scincidae. Of the 19 scincid type specimens in the Museum, the
majority were collected on the “Chevert” expedition (Macleay 18776), and
many still stand as holotypes. However, some of the material could not
be identified further than genus and status of such specimens is doubtful.
Three species described by Macleay could not be located in the collection.
The missing types are AHeteropus bicarinatus (18776), Tetradactylus
guttulatus (1885) and Hinulia picta (1885). Of these, the first is of the
most importance, as it stands as the holotype of Leiolopisma bicarinata
(Macleay) (see Loveridge, 1934). Tetradactylus guttulatus was ascribed to
the genus Lygosoma by Boulenger (1887), and Hinulia picta was synonymized
with Tiliqua gerrardii (Gray) by Mitchell (1950).
Fam. Gekkonidae. The gekkonid series in the Museum includes 10 types
described by Macleay (1877c), only one of which, Peripia dubia, is missing.
These types were examined in detail by Kluge (1963), who clarified their
systematic status as follows:
Diplodactylus annulatus = ? Phyllodactylus annulatus (Macleay) R487—488
Peripia papuensis = Hemidactylus frenatus Dumeril and Bibron R480-481
Peripia ornata = Lepidodactylus lugubris (Dumeril and Bibron) R484—485
Peripia longicaudis = Gehyra variegata (Dumeril and Bibron) R482
Peripia dubia Gehyra variegata (Dumeril and Bibron) Not present
Peripia marmorata = Gehyra baliola (Dumeril) R477
Peripia brevicaudis =Gehyra baliola (Dumeril) R475
Heteronota fasciata = Cyrtodactylus pelagicus (Girard) R176
Heteronota marmorata = Cyrtodactylus pelagicus (Girard) R178, 181-183
Heteronota eboracensis = Cyrtodactylus pelagicus (Girard) R207.
Fam. Agamidae. Five new agamid species were described by Macleay
(1877c, 1883), but only three types are still present in the Museum. Of
the two missing, one, Grammatophora jugularis (1877c), cannot be placed
further than the genus Amphibolurus (Boulenger 1885), and the other, Tiaris
longii (1877c), has been placed in synonomy with Gonyocephalus godeffroyi
(Boulenger 1885).
OrpER SERPENTES
Fam. Colubridae. Nine new colubrid snakes were collected on the
“Chevert” expedition (Macleay 1877a) and are all present in the Museum
today. Six additional species described by Macleay (1875, 1877e, 1883, 1884,
1888) are also listed. However, three of Macleay’s types are missing; viz.
Tropidonotus ater (1885), Fordonia variabilis (1877e) and Dendrophis
olivacea (1877e).
Fam. Elapidaec. Two new species of elapids were collected on the
“Chevert” expedition (Macleay 1877a). An additional eight type species
(Macleay 1878, 1884, 1885, 1887, 1888) are present in the Museum.
List or TYPES
The types are listed by families in current phylogenetic order.
An asterisk in the locality column indicates that the specimen(s) was
collected on the “Chevert” expedition.
References are given for the scientific name under which a type was
originally described and for the present name.
429
J. STANBURY
AND P.
HILL
GOLDMAN, L.
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SPECIMENS IN THE MACLEAY MUSEUM, II. AMPHIBIANS AND REPTILES
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SPECIMENS IN THE MACLEAY MUSEUM, II. AMPHIBIANS AND REPTILES
432
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SPECIMENS IN THE MACLEAY MUSEUM, II. AMPHIBIANS AND REPTILES
436
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Ayrore, 7
438 SPECIMENS IN THE MACLEAY MUSEUM, II. AMPHIBIANS AND REPTILES
Acknowledgement
The skeleton of this list was compiled by the previous curator of the
Macleay Museum, Mrs. J. M. E. Anderson, to whom we are indebted.
a
References
ANDERSON, J., 1965—MThe Macleay Museum at the University of Sydney. Aust. natural
History, 15: 47-51.
BouLENGER, G. A., 1882.—‘‘Catalogue of the Batrachia Salientia S. Ecaudata in the
Collection of the British Museum (Natural History).” 2nd ed. London. vii+127pp.
pis. i-ix.
: , 1885.—“‘Catalogue of the Lizards in the British Museum (Natural History).”
2nd ed. Taylor and Francis Ltd. London. 1: xii+436pp. 32 pls.
, 1887.—“‘Catalogue of the Lizards in the British Museum (Natural History).”
2nd ed. Tayor and Francis Ltd. London. III: xii+575pp. 40 pls.
CopLanpD, S. J., 1945.—Catalogue of the reptiles in the Macleay Museum. Part 1.
Sphenomorphus pardalis pardalis (Macleay) and Sphenomorphus nigricaudis
nigricaudis (Macleay). Proc. Linn. Soc. N.S.W.., 70 (5-6) : 291-311.
, 1946.—Catalogue of reptiles in the Macleay Museum. Part 2. Sphenomorphus
spaldingi (Macleay). Proc. Linn. Soc. N.S.W.,,71 (3-4) : 136-144.
Fry, D. B., 1913—A reexamination of Macleay’s New Guinea and Queensland frog
types. Mem. Queensland Museum, 2: 46-50.
Hann, D. E., 1962.—A list of the designated type specimens in the Macleay Museum.
Insecta. Roneoed list, University of Sydney.
Kuiver, A. G., 1963.—The systematic status of certain Australian and New Guinean
gekkonid lizards. Mem. Queensland Museum, 14: 177-86.
LOvERIDGE, A., 1934—Australian reptiles in the Museum of comparative Zoology,
Cambridge, Massachusetts. Bull. Mus. comp. Zool., 77: 243-383.
Macteay, W., 1875.—Notes on a new species of Dendrophis from Cleveland Bay. Proc.
Linn. Soc. N.S.W., 1 (1) : 15-16.
, 1877¢.—The Ophidians of the ‘‘Chevert’” expedition. Proc. Linn. Soc. N.S.W.,
m (ID) § SSeul
, 1877b.—The lizards of the ‘Chevert’” expedition. Proc. Linn. Soc. N.S.W.,
2 (1) : 60-69.
, 1877c.—The lizards of the ‘‘Chevert’” expedition. 2nd part. Proc. Linn. Soc.
N.S.W., 2 (1) : 97-104.
, 1877d.—The Batrachians of the ‘‘Chevert’” expedition. Proc. Linn. Soc. N.S.W.,
7) ((A)) 8 ileispilaxs):
, 1877e.—Notes on a collection of snakes from Port Darwin. Proc. Linn. Soc.
N.S.W., 2 (3) : 219-222.
, 1878.—On a new species of Hoplocephalus from Sutton Forest. Proc. Linn. Soc.
NSW. & (1) = 52-54:
, 1883.—Notes on some reptiles from the Herbert River, Queensland. Proc. LINN.
Soc. N.S.W., 8 (4) : 432-436.
, 1884—Census of Australian snakes, with description of two new species.
Proc. Linn. Soc. N.S.W. 9 (3) : 548-568.
, 1885.—On some reptiles from the Herbert River district, Queensland. Proc.
Linn. Soc. N.S.W., 10 (1) : 64-68.
, 1887.—On a new Hoplocephalus from the Gulf of Carpentaria. Proc. Linn.
Soc. N.S.W., 2 (2), 2nd ser. : 403-404.
, 1888.—Notes on some Ophidians from Kings Sound, North-west Australia.
Proc. Linn. Soc. N.S.W., 3 (2), 2nd ser.: 416-419.
MAcMILLAN, D. S. 1957—“A Squatter went to Sea.’ Currawong Pub. Co. Sydney.
MITCHELL, FrAnciIs J., 1950—A brief revision of the four-fingered members of the
genus Leiolopisma (ULacertilia). Rec. S. Aust. Museum.11: 75-90.
STANBuRY, P. J., 1968.—Type specimens in the Macleay Museum. 1. Fishes. Proc. Linn.
Soc. N.S.W. In press.
NOTES ON VITTADINIA TRILOBA sens. lat. (COMPOSITAE)
Nancy T. BuRBIDGE
Division of Plant Industry, O.S.1.R.0., Canberra, Australia
(Communicated by Dr. Joyce W. Vickery)
[Read 30th October, 1968]
Synopsis
Five taxa which have been included in Vittadinia triloba by some authors are
discussed. A new name is supplied, V. muelleri N. T. Burbidge, for a species believed
to represent part of Sonder’s Hurybiopsis hookeri var. angustifolia and also a new name,
V. blackii N. T. Burbidge is supplied for South Australian material formerly regarded
as V. tenuissima (Benth.) J. M. Black.
The genus Vittadinia A. Rich., Ess. Fl. Nouv.Zel. 250 (1832), is
distributed in Australia, New Guinea, New Caledonia and New Zealand, the
main group of species being.in the continental area. In the “Flora Australi:
ensis” Bentham took a broad view of species limits and it has long been
evident that a critical revision is required. In particular, the name V.
dustralis A. Rich. was applied to a range of material showing considerable
diversity and wide distribution in southern and eastern Australia. In more
recent Australian botanical literature this name has been replaced with
V. triloba Gaud. Preliminary study has indicated that some clarification of
this assemblage is possible and, pending a more intensive treatment, five
component taxa are considered below.
It has been found that achene characters are of diagnostic significance.
The outline is usually spathulate to oblanceolate or cuneate and there is
Some asymmetry which varies with position in the head, but the prominence
of the ribs on the flattened sides, as well as the nature and arrangement of the
hairs, varies from species to species.
V. TRILoBA- (Gaud). DC., Prodr. 5: 281 (1836); Brachycome triloba
Gaud., Bot., Freye. Voy. 467 (1830); V. australis A. Rich. sensu Bentham,
Fl. Austral. 3: 490 (1866); Hurybiopsis scabrida J. D. Hook., Lond. J. Bot.
6: 110 (1847); EH. hookeri F. Muell. ex Sond. var. scabra Sond., Linnaea
25: 454 (1853); V. scabra DC. sensu J. D. Hook., Fl. Tasm. 1: 181 (1856)
non DC. (1836).
The name V. australis A. Rich., to which Bentham referred the Australian
material, was based on a New Zealand plant in which the ligulate florets are
white, the inner involucral bracts about 5 mm. long and the trilobed leaves
are glandular-pubescent with ciliate margins. Since the Australian plants
have violet-coloured ligules, the inner involucral bracts are at least 6 mm.
long and the vestiture of the leaves is different, separation seems justified.
The type of Gaudichaud’s species was collected in the Port Jackson area.
The plants are scabrid with spreading multicellular hairs on stems and
leaves and also up the mid-line of the acuminate scales of the involucres; the
hairy achenes are narrowly turbinate-spathulate, ribbed and slightly flattened.
The distribution appears to be mainly in New South Wales, Victoria and
Tasmania. See Fig. 1 A (achene).
V. cUNEATA DC., Prodmvse2el 9(836)> a. D Hook., Fl. Tasmie obs 182
(1856); Hurybiopsis gracilis J. D. Hook., Lond. J. Bot. 6: 110 (1847): F.
PROCEEDINGS OF THE LINNEAN Society oF New SouTH WALES. Von. 93, Part 3
44() NOTES ON VITTADINIA TRILOBA
hookeri F. Muell. ex Sond. var. incana Sond., Linnaea 25: 454 (1853); V.
triloba var. lanuginosa J. M. Black, Trans. & Proc. Roy. Soc. S. Aust. 52: 229
(1928).
This species is distinguished by the woolly vestiture, especially on the
stems, the achenes are more flattened and more hairy than in V. triloba and
their narrow bases are clothed with appressed hairs. It is found in eastern
New South Wales, Victoria, south eastern parts of South Australia and in
Tasmania. See Fig. 1 B (achene) B, (bifid hairs).
V. MUELLERI, nom. nov.
Eurybiopsis hookeri F. Muell. et Sond. var. angustifolia Sond., Linnaea
25: 454 (1538) quoad “Van Diemensland (Stuart)”’.
Holotype—“Rockbank” adjoining Black Mountain Station, Wulgul-
merang, N. E. Gippsland, alt. 2800 feet approx., J. H. Willis, 27.xi.1962
(MEL 30018).
Plantae ascendentes, minute glanduloso-pubescentes. Folia angusta,
conduplicata, 1-4 cm. longa, integer vel lobis lateralibus, lobi angusti, divari-
cati, marginibus sparse ciliatis. Bracteae involucrales minute glanduloso-
pubescentes vel minute tuberculatae, marginibus membranaceis, apicibus
obtusis, ciliolatis. Achaenia striata, dimidio inferiore villis appressis, dimidio
Superiore pubescentia praeter marginibus, villis clavatis.
The plants are tufted, the minutely glandular-pubescent stems ascendent
with the narrow, more or less conduplicate leaves entire or with narrow
spreading lobes slightly above the middle, but leaves almost lacking from the
peduncles. Inner bracts of the involucres obtuse, without long hairs though
minutely glandular-pubescent or the surface appearing minutely tuberculate,
the margins ciliolate towards the apices. Achenes flattened, cuneate-
spathulate, the thickened margins glabrous, the lower portion with appressed
hairs grading into slender clavate (sometimes minutely bifid) hairs spreading
from between the ribs of the upper part, the ribs often inconspicuous on
immature fruits. Distribution widespread in eastern New South Wales,
Victoria and Tasmania. See Fig. 1 D (achene) D, (hairs).
New South Wales: 10 miles W. of Yarrowyck, New England, R. W.
Jessup & M. Gray 1794, 17.x.1952 (CANB); 2 miles N. of Dumaresq, R. W.
Jessup & M. Gray 1794, 17.x.1952 (CANB); Chiswick, 10 miles S. of
Armidale, R. W. Jessup & M. Gray 1750, 23.x.1952 (CANB); Cherry Hill,
Armidale district, R. Roe R548, 16.11.1945 (CANB) ; Kentucky, R. Roe R569,
21.11.1945 (CANB); 8 miles S. of Cessnock, R. Story 6708, 2.x.1959 (CANB) ;
Fairfield, O. D. Evans, 3.xii.1929 (CANB); Concord, O. D. Evans, 4.11.1927
(CANB) ; Mt. Jerrabomberra, S. of Queanbeyan, N. T. Burbidge 6696, 6.xi.1960
(CANB); Australian Capital Territory: near Burbong Village, Molonglo
River, P. J. Darbyshire 541, 21.xii.1961 (CANB, N.S.W.); Turner, Canberra,
R. Pullen, 1267, 21.11.1959 (CANB) ; Black Mountain, C. W. E. Moore, 5.x.1945 ;
also W. Hartley, 12.iv.1944 and R. Pullen 2061, 23.11.1960 (all CANB) ;
Kambah-Tharwa road, 3 miles past Kambah turn-off, M. Gray 3582, 18.xii.1958
(CANB); Michelago, New South Wales, H. S. McKee 7487, 23.x.1960
(CANB); Victoria: Heathcote, J. H. Willis, 7.xi.1961 (MEL 30018) ;
Tasmania: Mt. Nelson, C. E. Lord, January 1930 (CANB) ; Tasmania, Stuart
(MEL 30015).
Sonder listed two specimens under the variety angustifolia, one from
Van Diemensland and the other from Holdfast Bay (South Australia)
collected by Mueller in May. At Melbourne there is a Mueller specimen of
1851 from the Sonder Herbarium which is believed to represent the second.
NANCY ‘. BURBIDGH 441
No Stuart material that might be definitely associated as having been
examined by Sonder has been located. There is however a specimen on which
the label reads: “Kurybiopsis hookeri ferd MIll. var. laciniata, Tasmaniae
Stuart” in Mueller’s handwriting. Until proved otherwise this may be part
of the specimen seen by Sonder since it does, in fact, agree more closely
with his description than does the plant from Holdfast Bay. Sonder’s text
reads: “foliis . .. glabris vel subtus setulosis marginibus ciliolatis, inferiore
plerumque incisodentatis . .. achaenia puberulis.”
In the opinion of the writer the two specimens represent distinct species,
one described here and the other agreeing with material wrongly referred to
V. tenuissima (Benth.) J. M. Black and discussed below.
V. TENUISSIMA (Benth.) J. M. Black, Trans. & Proc. Roy. Soc. 8. Aust.
52: 229 (1928) quoad comb., descr. excl.; V. australis A. Rich. var. tenwissima
Benth., Fl. Austral. 3: 491 (1866).
ZB
— Ss
=
==
SS
SSS
neers
far
——— Aaa
= = = SS eee
= = Swe ey ete
SSsS50 Sane
SSE ee
pc
Fig. 1. A. Vittadinia triloba: achene. (from “Glenfield, O. D. Evans, 27.vii.1928.’).
B. V. cuneata: achene; B, hair. (from ‘Jerilderie, N.S.W., E. D’Arnay 388’). C.
V. tenuissima: achene; C, hair. (from “Grose Vale, Carne, N.S.W. 101651.”). D. V.
mueller: achene; D, hairs from upper part of achene. (from “Tharwa road Kambah,
A.C.T., Gray 3582.”) E. V. blackii achene; EH, hair. (from holotype.)
Bentham quoted Port Jackson specimens collected by Robert Brown
and by Woolls and a Mueller specimen from Burnett River (Queensland).
Judging by photographs of the first and third of these and by specimens in
the New South Wales Herbarium, the leaves are extremely slender and
almost or quite glabrous though the stems, which have a pronounced tendency
to branch corymbosely in the upper half above a simple base, are minutely ~
hairy. As noted by Bentham, the heads are small, the involucres being only
4—5 mm. long and the achenes 3-5-4:5 mm. long. This species is apparently
distributed from south east Queensland through the Northern Tablelands
of New South Wales and south through coastal districts to near the Victorian
border. Fig. 1 C (achene) C; (hair).
Though Black based his combination on Bentham’s varietal name his
description does not fit the type material. He states “involucrum 7-8 mm.
longum .. . achaenia 4-5 mm. longa puberula utrinque circiter 6—costata .. .”
K
449 NOTES ON VITTADINIA TRILOBA
He also mentions the ligules as numbering 15-20 which is higher than in the
eastern Australian specimens examined. It is thus evident that though his
combination is the correct name for the species typified as Bentham’s var.
tenuissima, his description covers a different species. This latter is discussed
below.
V. BLACKII, nom. nov.
V. tenuissima (Benth.) J. M. Black quoad descr., basion. excl.; Hury-
biopsis hookeri F¥. Muell. et Sond. var. angustifolia Sond., Linnaea 25: 454
(1853) quoad “Holdfast Bay, Mai”.
Holotype.—Old Stockade Hill, Northfield, ca. 10 km. N.N.E. of Adelaide,
South Australia, D. N. Kraehenbuehl 1538, 5.iii.1960 (AD 96422044).
Plantae minute glanduloso-pubescentes, villis septatis sparsim ornatae.
Folia lineari-teretia vel lineari-conduplicata, 1-1-5 (-3) cm. longa. Bracteae
involucrales minute glanduloso-pubescentes, marginibus membranaceis,
apicibus ciliatis. Achenia striata, dimidio inferiori glabra villis appressis
infra exceptis, dimidio superiori minute pubescentia, villis brevibus crassis
clavatis etiam ornata.
Plants subshrubby, 10-380 cm. high, older stems more or less decumbent
and woody, stems much branched, bearing septate hairs mixed with minute
glandular pubescence. Leaves linear, conduplicate but usually so narrow as
to appear terete and channelled above; mostly 1-1-5 cm. long but occasionally
longer with sparse scattered hairs and few minute glandular ones, the surface
more or less glistening. All stems terminating in solitary heads, involucres
7-9 mm. long, bracts with green centres bordered with minute glandular
hairs, a few longer hairs sometimes present, margins membranous and ciliolate
towards the acuminate apex. Achenes shorter than innermost bracts,
flattened, narrow cuneate, deeply ribbed, with scanty appressed hairs at
base but otherwise glabrous in lower half, the upper half with projecting
short clavate hairs, margins glabrous, pappus bristles very numerous, smooth
towards base but barbellate above. Fig. 1 E (achene) E; (hair). Distribution
mainly restricted to South Australia but also recorded from western New
South Wales and north eastern Victoria with one record from Western
Australia.
This species can be separated from V. muelleri by the septate hairs on
the stems, the scattered hairs of the leaves and by the deeply ribbed achenes
with fewer and short clavate hairs attached to the ribs rather than between
them.
Western Australia: Halfway between Mt. Ragged and Victoria Spring,
Miss S. Brooks, 1886 (MEL 30012); South Australia: Birksgate, M. Koch,
Sept. 1902 (NSW 101654) ; Lake Eyre Basin, Schomburgk (AD 96826384) ;
Hambidge Flora and Fauna Reserve (ca. 140 km. N. of Port Lincoln) Eyre
Peninsula, C. R. A. Alcock 1103, 10.x.1966 (AD 96711206); Thrington on
Thrington-Moonta road, Upper Yorke Peninsula, B. Copley 163, 27.iii.1966
(AD 96622021); Maitland, Yorke Peninsula, J. M. Black, April 1917
(AD 96826385) ; ca. 2 km. S. of Hamilton on road to Kapunda, Mt. Lofty
Range, Hj. Eichler 12074, 7.xii.1955 (AD 95902036) ; Freeling Cemetery, ca.
55 km. N.N.E. of Adelaide, D. N. Kraehenbuehl 1505, 18.ix.1965 (AD
96724004) ; Adelaide Plains near Adelaide, J. M. Black, April 1917 (AD
9682385) ; North bank of Dry Creek, east of Yatala Prison Farm, Adelaide
Plains, D. N. Kraenhenbuehl 465, 13.ix.1961 (AD 96426228); Brighton,
J. M. Black, 18.ix.1904 (AD 96826386); Echunga district, R. F. Parsons
212, 27.x.1961 (AD 96348243): Murray Bridge, J. H. Maiden, January 1907
NANCY T. BURBIDGE 443
(NSW 101655) ; Sandergrove, O. IX. Menzel, Oct. 1896 (AD 96826380) ; Port
Elliot, Fleurieo Peninsula, J. B. Cleland, 25.13.1925 (AD 9682383); sine loc.,
Behr, 10.11.1845 (MEL 30009) (type of Aster behrii?) New South Wales:
Interior, Behr (MEL 380016) ; Victoria: Murray Desert, Behr (MEL 30011) ;
Pine Plains, Wimmera, Behr 215 (MEL 30017).
Key to species discussed
1. Leaves spathulate, oblanceolate or cuneate, entire or 3—-lobed at the apex. Achenes
5-6 mm. long, narrowly oblanceolate, vestiture on both sides and margins consisting
of slender hairs with bifid apices above underlying glandular pubescence.
Plants clothed with soft woolly hairs; basal part of achene with dense appressed
hairs, the hairs of the upper part spreading and obscuring the ribs:
cuneata
bho
2a. Plants scabrid with spreading septate hairs; lower part of achene almost glabrous
except for short appressed hairs at base, upper part prominently ribbed, slightly
flattened but turgid, hairs spreading:
triloba
la. Leaves filiform, linear or narrowly elliptical-oblanceolate, entire or sometimes
with a pair of spreading narrow lobes near or above the middle; achenes narrowly
cuneate or spathulate-cuneate, margins glabrous or almost so.
3. Leaves filiform, almost or quite glabrous; involucres 4-5 mm. long, achenes
35-4 mm. long; lower half glabrous apart from short appressed hairs at base,
upper half with sparse slender spreading hairs with bifid apices: glandular
pubescence lacking:
tenuissima
3a. Leaves sparsely hairy, hairs septate; involucres 6-9 mm. long; achenes 4-5-5 mm.
long, minute glandular pubescence present below hairs.
4. Leaves with scanty hairs on margins and on midrib of lower surface; lower part
of achene clothed with slender appressed bifid-tipped hairs which grade into
slender obtuse or bifid-tipped clavate hairs growing between the ribs which are
often inconspicuous before maturity:
muelleri
4a. Leaves with scattered hairs; achenes with appressed slender hairs at base of
glabrous lower half, deeply striate above with 6-7 prominent ribs on each side
even when young, the ribs bearing short clavate hairs:
blackii
A REVIEW OF THE FAMILY:AGNESIIDAE HUNTSMAN 1912;
WITH PARTICULAR REFERENCE TO AGNESIA GLACIATA
MICHAELSEN, 1898
Parricia Korr
Zoology Department, University of Queensland
[Read 27th November, 1968]
Synopsis
Known species of the family Agnesiidae are reviewed and their relationships
are clarified. Particular attention has been given to the genus Agnesia and the
synonymy and distribution of Agnesia glaciata. New occurrences are recorded for
Agnesia glaciata and Adagnesia opaca.
Similar modifications of body musculature to operate specialized closing mechanisms
are demonstrated in each genus of the family. The development of this protective
closing mechanism is associated with the extent to which the test is made brittle
and rigid with encrusting sand thus preventing the general contraction of the body
as a defence mechanism. The three known genera are closely related and distinguished
by branchial sac modifications from Caenagnesia through Adagnesia to Agnesia.
Adagnesia opaca and Agnesia glaciata are the most specialized species. The family
appears to be an ancient one and records indicate relict populations of all species.
INTRODUCTION
The family Agnesiidae of the Suborder Phlebobranchia contains a
limited number of closely related and highly specialized genera. Records of
the family are not common, although often large numbers of individuals
are taken together.
The following genera are known:
(1) Caenagnesia Arnbiick, 1938, is known only from Antarctica and
is represented by two species.
(2) Agnesia Huntsman, 1912, is represented by one species from the
north Pacific; a second species extends from California to Tierra del Fuego
and the Antarctic Peninsula, North Island, New Zealand, South Africa,
Moreton Bay, Queensland, and Japan; a third species is known from abyssal
depths of the north Atlantic.
(3) Adagnesia Kott, 1963, of which 2 species are aoe one from a
Single specimen off Macquarie Island, and one from a limited area of the
Australian coast.
In the present work the inter-relationships of species of this family are
discussed, especially in regard to the increasing specialization of body
musculature.
Family AcnesiipAn Huntsman, 1912
Gut on the left side of the branchial sac; internal longitudinal vessels
reduced to papillae on the transverse vessels; stigmata spiral; branchial
PROCEEDINGS OF THE LINNEAN SocreTy oF NEw SoutTH WALES, VOL. 93, Part 3
PATRICIA KOTT L415
tentacles arranged in 4 concentric circles; the border of the branchial and
atrial apertures produced into 6 and 7 pronounced lobes respectively: muscle
bands reduced in length, often considerably.
Throughout the family Agnesiidae there is an increasing reduction in
the numbers of papillae on the transverse vessels; reduction in numbers of
infundibula; and a reduction in the number of transverse vessels present.
The musculature becomes highly specialized and there is a progressive
reduction in numbers and length of muscle bands throughout the family.
Despite the specialized nature of body musculature in the Agnesiidae,
the homologues of the cionid musculature can be traced. In Ciona intestinalis,
the most primitive species known, the external layer of musculature is
represented by longitudinal bands and the internal layer consists of circular
fibres. Only on the siphons, anterior to the tentacular band, does the
circular muscle layer become superficial to the longitudinal bands. The
internal transverse musculature, present more posteriorly in Agnesiidae,
represents the inner layer of circular fibres in Ciona. While the anterior
and superficial transverse bands represent the continuation of the circular
bands which in Ciona are confined to the siphons. The inner circular bands
associated with the tentacular ring in Agnesiidae are also a vestige of the
inner circular layer of Ciona. The most significant departure from the cionid
condition observed in the musculature of Agnesiidae is the reduction in length
and number of longitudinal muscle bands; and the interruption of circular
bands to form shorter transverse bands confined to the dorsal and ventral
borders of the body. True circular bands are, in Agnesiidae, confined to the
siphons. On the rest of the body the circular musculature is interrupted
first laterally (Caenagnesia spp.) and then also in the median dorsal and
ventral lines (Agnesia spp. and Adagnesia spp.).
The shortened muscle bands of Agnesiidae are associated with increasing
rigidity of the test due to encrustation with sand. The body is consequently
less contractile and the functions of the muscles become more specialized.
In Caenagnesia bocki, Agnesia glaciata and Adagnesia opaca the shortened
muscle bands pull lips or folds of test, which is generally rigid with sand,
across the apertures to form a closing mechanism. This undoubtedly serves
as a protective device for these non-contractile species existing in a vulnerable
sublittoral locality.
The subfamily Rhodosomatinae of the family Corellidae, also of the
suborder Phlebobranchia, contains monotypic Rhodosoma turcicum (Savigny),
the only species outside the present family which exhibits a similar closing
mechanism protecting the apertures. In Rhodosoma this is less symmetrical
but more conspicuous than in the Agnesiidae and a fold involving the body
wall and test is developed only on the right side of the apertures to form
a lid. This is operated by the transverse muscle bands at the base of the
siphons, particularly those across the mid line between the apertures, which
extend out into the fold (Kott 1952). The mechanism is similar in Agnesiidae
where the circular muscles surrounding the base of the siphons are interrupted ~
laterally and operate across the dorsal midline from the base of the test fold
on either side of the apertures. The mechanism appears to develop
independently in each genus of the family. Closing lips are formed in
Caenagnesia bocki and are present but not so well developed in Adagnesia
antarctica which appears to have been derived from Caenagnesia. In
Adagnesia opaca the closing mechanism achieves its greatest development
and in Agnesia, a genus which probably evolved from a primitive Adagnesia
446 A REVIEW OF THE FAMILY AGNESIIDAE HUNTSMAN
sp. a gradual specialization of musculature to operate closing lips is observed
within a single species, Agnesia glaciata, where, in its most specialized form,
it closely resembles the mechanism found in Adagnesia opaca.
Genus CAENAGNESIA Arnbick, 1938
Traces of longitudinal vessels remain as bifid papillae on the transverse
vessels of the branchial sac. Dorsal lamina retains the primitive condition
of a plain edged membrane. Primary transverse vessels bearing papillae
are present between each row of infundibula. Branchial papillae are more
numerous than the number of infundibula in each row and more than a
single papilla corresponds to each infundibulum. There are at least 12
primary transverse vessels present.
CAENAGNESIA SCHMITTI Kott
(Text Fig. 1)
Caenagnesia schmitti Kott, 1969, p. 94.
Specimens examined.—U.S. National Museum: South Shetlands,
“Eltanin”, St. 428, 662-1120 m; coll. W. Schmitt, St. 66/63, 62 m. Victoria
Land, S. W. Robertson Bay, 400 m (Holotype USNM 11968). (Single
specimens from each station.)
Remarks.—The body is cylindrical, the test is thin and the body wall
highly contractile. The apertures are on short cylindrical siphons which are
furrowed along their length. The longitudinal muscle bands are especially
numerous (about 50 on each side) and extend the whole length of the body on
the right and as far as the gut loop on the left. Contraction of these muscles
tends to draw the gut loop up along the branchial sac; however, in relaxed
specimens the gut loop appears to lie behind the branchial sac reminiscent
of the situation in Ciona intestinalis. Circular muscles are present around
the siphons, external to the longitudinal muscles and numerous fine circular
bands are present internally, associated with the 4 circles of branchial
tentacles. Ventrally a superficial layer of transverse muscles is present
anteriorly and is continuous across the endostyle. About 5 of the most
posterior bands of this series overlap, in the middle of the body, with an
inner transverse series present posteriorly on either side of the endostyle
but not continuous across it. Dorsally a similar series of superficial trans-
verse muscles is continuous across the dorsal line in the anterior two thirds
of the body. These overlap with approximately the 5 most anterior bands
of an inner transverse series which are interrupted across the dorsal line.
The anterior superficial transverse bands, continuous across the dorsal
surface, extend across the distal part of the rectum; while the inner and
more posterior bands, interrupted across the dorsal surface, allow for some
expansion of the stomach and proximal part of the rectum. Ventral trans-
- verse muscles are longer than the dorsal bands and neither extend across
the sides of the body. In the branchial sac the number of rows of infundibula
(60) and the number of infundibula in each row (25) are especially numerous
for species of this family. The number of papillae on the transverse vessels
are also numerous, with especially long biramous arms. There is a simple
dorsal lamina as in Ciona intestinalis and other families of Phlebobranchia.
Occurrence.—Antarctica, probably circum-antarctic continent 74 m to
1000 m (Kott 1969).
PATRICIA KOTT 447
Discussion.—The shape of the body, the furrowed external siphons, the
thin test and the large numbers and length of the longitudinal muscles are
unspecialized and reminiscent of Ciona intestinalis. The branchial sac also
shows primitive affinities, as the number of rows of stigmata and the number
BE 1.0cm
EZ
- &
Sa a
= Re
Lf
Ath
ct
u
1.0c¢m
1.0 cm
Text-figure 1-7 (Semi-diagrammatic showing body musculature) 1. Caenagnesia schmitti
(Relay Bay, Antarctica). 2. Caenagnesia bocki (South Shetlands, Antarctica). 3.
Agnesia septentrionalis (St. Georges Sound, Probilof Is.). 4. Agnesia glaciata (Corona
del Mar, California). 5. Agnesia glaciata (South Shetlands, Antarctica). 6. Agnesia
glaciata (Antarctic Peninsular, Antarctica). 7. Agnesia glaciata (Moreton Bay,
Queensland).
448 A REVIEW OF THE FAMILY AGNESIIDAE HUNTSMAN
of papillae per row is reduced in more specialized forms; and the retention
of long biramous arms on the papillae suggests that the reduction of the
longitudinal vessels has not proceeded as far as in other species with smaller
papillae.
CAENAGNESIA BOCKI Arnbick
(Text Fig. 2)
Caenagnesia bocki Arnbiick, 1938, p. 41. Van Name 1945, p. 202. Millar
1960, p. 94. Kott 1969, p. 96. Agnesia complicata Kott 1954, p. 151.
Specimens examined—uvU.S. National Museum: Weddell Sea, “West
Wind”, St. 4, 796 m; “Edisto”, St. TR6, 394 m; South Shetlands, coll. W.
Schmitt, St. 9/63, 57 m; “Eltanin”, St. 437, 267-311 m. Australian Museum:
BANZARE Collection: Enderby Land, 220 m. (Single specimens.)
Remarks.—tThe species is dorso-ventrally flattened and the almost sessile
apertures are fairly close together on the upper surface. Folds of test are
formed along each side of the apertures into which the body wall projects.
These folds may meet along the median line, thus covering and protecting
the apertures. The test is firm and transparent and is sometimes brittle
with sand. Due to the dorso-ventral flattening of the body the gut appears
to lie across the posterior surface only slightly to the left. The longitudinal
muscles are numerous (about 20) on each side. They extend the whole length
of the body on the right but only to the gut loop on the left. There are
circular muscles on the siphons external to the longitudinal muscle. Internally
there are about 11 circular muscles associated with the 4 circles of tentacles.
In a continuous series with the external circular siphonal muscle bands there
are transverse bands across the dorsal border of the body in the anterior
two thirds. Similar transverse muscles are present in the anterior one third
of the body ventrally. The most anterior of these transverse muscle bands
are inserted into the body wall at the base of the protective folds of test.
Overlapping with the external transverse muscles there is a layer of internal
transverse musculature across the dorsal border on the inner surface of the
proximal part of the gut loop; and across the endostyle ventrally. More
diffuse musculature across the posterior end of the body is gathered into
bands which are inserted into the area enclosed by the pole of the gut loop
where they are obscured by gonad lobes. All transverse muscles are interrupted
across the sides of the body. The protective closing mechanism operates by
the contraction of the external transverse muscle bands anteriorly which
pull the lips together; meanwhile the siphonal musculature closes the
apertures.
The branchial sac in this species is reduced from the condition found in
C. schmitti. There are primarily 12 rows of 13 to 14 spiral infundibula. With
growth the number of rows multiplies and a specimen of 30 mm (Kott 1954)
has 24 rows of about 17 spirals. Biramous papillae with long arms are
present on the transverse vessels between each row of infundibula. There
are about 3 corresponding to each spiral.
Occurrence.—South Shetlands, Antarctic Peninsula (Arnback 1938, Millar
1960, Kott 1969); South Georgia (Millar 1960) ; Weddell Sea (Kott 1969) ;
Enderby Land (Kott 1954); at 57-800 m.
Discussion.—This species is distinguished from C. schmitti especially by
the reduced numbers of branchial papillae, transverse vessels and infundibula,
PATRICIA KOTT 449
and by the protective lips which cover the apertures. The reduced number and
length of longitudinal muscle bands and the reduction in number of internal
circular muscles associated with the tentacular sphincter is probably related
to the development of a closing mechanism associated with the loss of a
flexible body. The internal muscle bands are continuous posteriorly and
dorsally across the mesial surface of the gut loop beneath the retropharyngeal
groove that extends from the oesophageal opening at the base of the dorsal
lamina to the posterior end of the endostyle. In Aplousobranchia the mascula-
ture of the body wall extends outside the gut loop, generally to the posterior
end of the body. It has been suggested (see Kott 1969, p. 190) that the
position of the gut loop on the side of the branchial sac could have resulted
from a backwards extension of the branchial sac to the right or left of the
gut loop. The muscle bands would, in this case remain outside the gut
loop. The presence of the internal transverse muscle bands on the mesial
surface of the gut loop in the present species suggests that, at least in
this family, the gut has been.-drawn up on the left side of the branchial
sac by relative shortening of the longitudinal muscle bands, leaving the
inner transverse muscles across the posterior end of the body.
Genus AgnestA Michaelsen, 1898
Simple undivided flat triangular papillae present on transverse vessels.
Dorsal lamina absent. Area of flat unperforated membrane crossed by
primary transverse vessels present along mid dorsal line of branchial sac.
Enlarged triangular papillae on the primary transverse vessels to the left of
the dorsal line may correspond to dorsal languets. The number of infundibula
present in each row always exceeds the number of papillae present on the
transverse vessels. Four primary transverse vessels present (crossing the
mid dorsal line).
AGNESIA SEPTENTRIONALIS Huntsman
(Text Fig. 3)
Agnesia septentrionalis Huntsman, 1912, p. 118; 1912a, p. 106. Van
Name, -1945, p. 200. Agnesia beringia Ritter, 1913, p. 493.
Specimens examined.—uvU.S. National Museum (Cat. No. 10633): coll. G.
W. Hanna, St. George Island, Alaska (5 specimens). American Museum of
Natural History (Cat. No. 1896) : coll. G. Hanna, St. Georges Sound, Pribilof
Islands, 74 m (8 specimens).
Remarks.—Rounded oval body from 0-5 to 1:5 cm long, usually supported
by a delicate short stalk from the posterior end of the body. The stalk
may be 1 cm long on a specimen of 0-8 cm but is generally much shorter
than this. The test is thin and glassy, with few hairs and adherent sand
erains. The apertures protrude on short rounded siphons, the branchial
aperture anterior, and the atrial aperture antero-dorsal. There are 4 rows
of equally long and closely placed branchial tentacles and posterior to these,
at the base of the siphon, a ring of about 6 circular muscles which may
coalesce into a single wide band in the rim of the tentacular velum. About
30 longitudinal muscle bands radiate from each siphon for only a very short
distance. Externally each siphon has about 22 circular muscle bands. On
the atrial siphon these extend well down the dorsal surface to cross the
rectum. Short transverse muscle bands are arranged in single series on
either side of the dorsal and ventral median lines, beneath the longitudinal
_ muscles, and, especially on the dorsal surface, these extend anteriorly beneath
450, A REVIEW OF THE FAMILY AGNESIIDAE HUNTSMAN
the circular siphonal musculature. In this species the branchial sac is well
developed. There are 4 primary transverse vessels, bearing large flattened
tongue-like papillae. Between successive primary transverse vessels 4 rows of
22 infundibula develop from double rows of 11 primary spirals. Secondary
transverse vessels develop about 7 papillae corresponding to those on the
primary vessels. Intermediate transverse vessels of a third order separating
the double rows of infundibula between the primary and secondary vessels
also develop papillae but these are generally incomplete. Each infundibulum
develops a maximum of 4 to 5 coils.
‘Occurrence.—Bering Sea (Ritter 1918); British Columbia (Huntsman
1912, 1912a); Alaska, Pribilof Islands (Ritter 1913); at 27 to 78 m.
Discussion.—Differences between this species and A. glaciata involve the
development of the body musculature which is less complex in the present
species and the siphonal muscles in particular are not modified to operate
a special closing mechanism. Apertures close by simple sphincter muscles
around the siphons and these circular muscles are never interrupted. Similarly
the branchial tentacles are more conspicuous in the present species and the
inner circular muscles at the base of the branchial siphon associated
with a velum are better developed. Development of the branchial sac depends,
in the present species, on a proliferation of primary infundibula. In A.
glaciata growth is accompanied by increase in the number of coils of each
infundibulum.
AGNESIA GLACIATA Michaelsen
(Text Figs 4-7)
Agnesia glaciata Michaelsen, 1898, p. 370; Van Name, 1945, p. 200;
1900, p. 76; 1907, p. 75; Millar, 1960, p. 92; Kott 1969, p. 97. ‘Agnesia krauset
Michaelsen 1912, p. 181. Agnesia capensis Millar 1955, p. 191. Agnesia
himeboja Oka 1915, p. 1. Agnesia sabulosa Oka 1929, p. 152. Agnesia septen-
trionalis; Van Name, 1945, p. 201, Part (specimens from Newport Harbour,
Southern California). .
Specimens examined.—U.S8. National Museum: Antarctic Peninsula, coll.
W. Schmitt, St. 27/63, 75 m; 66/63, 74-92 m (single specimens); South
Shetlands, coll. W. Schmitt, St. 64/63, 86 m (2 specimens). American Museum
of Natural History (catalogue Nos. 1570, 1571, 1572, identified as A.
septentrionalis by Van Name 1945): coll. MacGinitie; Corona del Mar
California. Queensland Museum (Registration No. G5214): coll. W. Stephen-
son, Moreton Bay, Queensland 27°14’50’S, 153°18’00”E, 23 m; 27°16’20’S
153°20’50°E, 24 m (numerous specimens on muddy sand).
Remarks.—Mature specimens vary in size from less than 1-0 em in
diameter to 4 em long. They are generally more or less rectangular and
contracted specimens may be dorso-ventrally or laterally flattened. Both
siphons are sessile and are present on the anterior surface. Variations in
morphological characters occur in association with increasing size or
increasing rigidity of test due to sand encrustation. Without sand the test
is thin and semitransparent and may have fine hairs especially from the
posterior end.
In large specimens from the South Shetlands (coll. W. Schmitt, 3 to
4 cm long) the test is free of sand and is very delicate posteriorly while
the anterior half of the body is thickened, except for an area immediately
surrounding the siphons. On contraction, the anterior thickened portion of
the test forms a protective dome over the dorso-ventrally contracted body
PATRICIA KOTT 15]
and the siphons are drawn down into, and are protected by, the surrounding
thicker test. Smaller specimens from California (Newport Harbour coll,
MacGinitie, 1:0 em) and from Patagonia (A. krausei Michaelsen 1898, 0-9 em)
have the test thickened in a similar way although there is some sand adhering.
Contraction causes a withdrawal of the apertures which are protected by the
sandy ridge of thickened test surrounding them. Specimens from Moreton
Bay (1:5 cm), from Tierra del Fuego (Michaelsen 1898, 1:8 em) and the
specimen from the Antarctic Peninsula (Kott 1969, 2 em) are heavily
encrusted with sand causing the test to be rigid. Here the area of test
surrounding the apertures is extended into a fold involving the body wall.
On contraction the body is laterally flattened and the fold, especially from the
right side, closes over the apertures.
The body musculature basically consists of about 20 superficial circular
muscles around the branchial siphon and usually more associated with the
atrial siphon. The basal muscles on the atrial siphon are not continuous
around the anterior side of the siphon. These are referred to as posterior
atrial muscles and they extend well down the dorsal surface. There is only
a single circular band associated with the tentacular velum. From each
side of the siphons longitudinal bands radiate not more than half-way down
the body. There are more longitudinal bands frem the branchial than from
the atrial siphons. The proximal 8 to 10 circular bands around each aperture
have fibres joining the longitudinal bands; and the posterior atrial muscles
also have some fibres joining the distal extent of the longitudinal muscles.
In specimens from California there are about 20 circular bands around
the branchial siphon; about 15 circular bands around the atrial siphon; and
6 posterior atrial bands. There are 14 longitudinal bands from each side
of the branchial siphon and 8 from each side of the atrial siphon. In the
specimens from the South Shetlands there are 18 longitudinal bands from
either side of the branchial aperture; 15 from either side of the atrial aperture
and 6 to 15 posterior atrial bands. The circular bands vary from 20 to 50
on both siphons. Where a protective fold of test is developed (specimens
from Moreton Bay) the anterior 8 circular muscles of the branchial and atrial
siphons remain unmodified and remain associated with the proximal extent
of the longitudinal bands. However the next 4 to 7 circular bands break up
into fibres dorsally, ventrally and on both sides of each siphon in the fold
of the body wall associated with the test fold. These bands are not associated
with the longitudinal bands. Posteriorly to these, a further 10 muscle bands
on either side of the dorsal line are homologous with the posterior atrial
bands in other specimens and some fibres branch into the longitudinal bands;
5 similar bands on either side of the ventral line represent modified circular
bands from the base of the branchial siphon and are also associated with
the longitudinal bands. In these specimens there are 11 longitudinal bands
on either side of the branchial siphon and 6 longitudinal bands on either
side of the atrial siphon. In the larger specimen from the Antarctic
Peninsula 12 circular bands remain uninterrupted around the apertures and
the next 7 circular muscles are associated with the protective folds and break
into fibres on each side of the apertures. In this specimen these muscles
are continuous across the dorsal and ventral surfaces. This is a more
posterior group of muscles than those associated with the fold in the Moreton
Bay specimens and only 5 short bands remain posterior to the folds, on either
side of the dorsal line and associated with the longitudinal bands. In this
specimen there are 25 longitudinal bands per side.
452. A REVIEW OF THE FAMILY AGNESIIDAE HUNTSMAN
The data given above indicate that, with an increase in size of the body,
there is an increase in the number of longitudinal muscle bands and the
numbers of bands associated with the siphons. There is also considerable
variation in the relative numbers of bands utilized for different functions.
The longitudinal muscles are never involVed with the fold or ridge surrounding
the apertures and their function appears to be to draw the apertures down
into the body and to dorso-ventrally flatten the body. They are consequently
longer when the test is less rigid. The uninterrupted circular bands act as
siphonal sphincters. However, when the protective fold is developed, it is
the circular bands from the middle of the siphons which, together with a
varying number of posterior atrial muscles, break into fibres and operate the
protective folds of test. The number of circular bands which remain entire
in the anterior extent of the siphon affects the number of posterior atrial
muscles which become involved with the closing mechanism and seems to
depend on the extent to which the siphons are developed. Consequently,
in the large specimen from the Antarctic Peninsula where 12 muscle bands
operate as a sphincter, the posterior atrial bands contribute to the operation
of the closing mechanism leaving only a few of these below the folds. In
the Moreton Bay specimens 8 muscle bands operate as a sphincter, the
remainder of the circular siphonal muscles are modified to operate the
closing mechanism, and the posterior atrial muscles, interrupted across the
dorsal line, all remain behind the protective fold and probably contribute
to the lateral flattening of the body. Unmodified posterior atrial bands,
inserted into the body wall where it is associated with the thicker test around
the siphons, probably draw the atrial siphon towards the branchial siphon
and draw the test in more closely around both siphons.
In addition to the muscle bands described above short muscle bands
deep to the longitudinal bands are continuous around the dorsal and ventral
borders of both sides of the body. These muscles tend to flatten the body.
The dorsal tubercle is a simple slit and there is a tongue-like evagination
of the body wall projecting from the region of the dorsal gland. The branchial
sac has 6 double rows of 11 infundibula. Transverse vessels between each
double row bear triangular papillae which are enlarged to the left of dorsal
line. These transverse vessels are continuous over the dorsal membrane. A
dorsal lamina is not formed. Only in the specimen from the Antarctic
Peninsula are papillae developed also on the intermediate transverse vessels.
The latter do not cross the dorsal line. Stigmata form 8 to 10 spirals. These
are often interrupted in the vertical or horizontal axis and are crossed by
radial vessels.
Occurrence.—Antarctic Peninsula, South Shetlands (Kott 1969); Tierra
del Fuego (Michaelsen 1898); Patagonian Shelf (Michaelsen 1912, Millar
1960); California (Van Name.1945); New Zealand (Millar 1960); South
Africa (Millar 1955, 1960) ; Moreton Bay, Queensland (New records) ; Japan
(Oka 1915, 1929); at 23 and 115 m.
Discussion.—The tremendous variation in the disposition of the body wall
musculature together with the widely dispersed records suggests that more
than a single species is involved here. It is apparent however that the
muscles are disposed merely in response to the degree of rigidity of the test.
Where a heavy sand incrustation prohibits contraction of the anterior test
around the withdrawn siphons the test instead folds over the siphons and
the muscle bands break up into fibres in the folds. The numbers of muscle
bands which are modified to effect the various contractions required of the
body wall are immensely varied and the most posterior circular bands may
PATRICIA KOTT 453
either be involved in the protective fold or may remain superficial to the
longitudinal bands posterior to the fold. This may depend to some extent
on the size of the fold; or on the relative size of the body; or the development
of the siphons.
In younger specimens (A. capensis Millar, A. himeboja Oka, A. sabulosa
Oka, A. krausei Michaelsen), the apertures remain in the primitive position,
the branchial aperture anteriorly and the atrial aperture antero-dorsally and
the ridge of test protecting them has not developed. The interrupted muscles
bands behind the atrial siphon, the number of coils in each fundibulum, the
number of infundibula and transverse vessels and the single muscle band in
the tentacular velum all indicate that these specimens do fall within the range
of variation of the species.
The species is distinguished from Agnesia septentrionalis by the larger
number of spirals in each infundibulum; by the fact that at least some circular
muscles are not completely continuous around the siphons; and by single
muscle band associated with the tentacular velum.
AGNESIA DEPRESSA Millar
Agnesia depressa Millar, 1955a, p. 1.
Remarks.—Millav’s specimens (4) ranged from 0:8 to 1:1 cm in greatest
diameter and this species resembles other Agnesia spp. in the presence of
hair like processes from the test and in the numbers and arrangement of
transverse vessels and infundibula. The species is distinguished only by the
reduction of the branchial papillae and is apparently closely related to
Agnesia glaciata.
Occurrence.—Swedish Deep-sea Expedition Sta. 371; N 24°12’, W 63°23’
to N 24°28’, W 63°18’; 5850 to 5860 m (Millar 1955a).
Discussion.—It is remarkable that a species from this depth shows so little
deviation from other species. Apart from the longer test hairs it demonstrates
none of the usual modifications, especially of the branchial sac, which
are usually associated with abyssal species.
Genus ApacnestA Kott, 1963
Bifid papillae on the transverse vessels. Dorsal lamina absent. Area
of flat unperforated membrane crossed by primary transverse vessels present
along mid-dorsal line of branchial sac. Enlarged triangular papillae on the
primary transverse vessels to the left of the dorsal line may correspond te
dorsal languets. The number of infundibula always exceeds the numbers of
branchial papillae in each row. Always more than 4 primary transverse
vessels present.
ADAGNESIA ANTARCTICA Kott
Adagnesia antarctica Kott, 1969, p. 99.
Specimen examined.—U.S. National Museum: West of Macquarie Island,
“Eitanin”, St. 1418, 86-101 m. (Holtype U.S.N.M. 11966.)
Remarks.—The single available specimen is rounded and 1°55 cm in
diameter. The test is thin but brittle and encrusted with sand. The apertures
are surrounded by a rim of thickened test around the upper surface. There are
about 20 circular muscles around each siphon and 35 longitudinal muscles
on each side of the body extending only a short distance from the base
of the siphons. Short transverse muscle bands are present in rows around
the dorsal and ventral borders on either side of the body. In the branchial
454 A REVIEW OF THE FAMILY AGNESIIDAE HUNTSMAN
sac there are 7 double rows of spiral infundibula separated by 6 primary
transverse vessels. Biramous papillae with long arms are present on the
primary transverse vessels but there is never more than one of these papillae
corresponding to each spiral and often there are fewer. Along the dorsal
line a large single languet develops frons each primary transverse vessel, as in
Agnesia.
Occurrence.—Only the holotype from west of Macquarie Island is known.
Discussion.—The species is distinguished from Caenagnesia spp. by reduc-
tion in the numbers and length of longitudinal muscle bands, a reduction in the
length of transverse bands and their interruption over the mid line, and a
reduction in the numbers of circular siphonal muscle bands; a reduction in
the numbers of primary transverse vessels, infundibula and _ branchial
papillae; the loss of the dorsal lamina and the presence of enlarged dorsal
languets on the primary transverse vessels on the dorsal line. In the condition
of the dorsal lamina and numbers of infundibula in each row, the species
resembles Agnesia spp. However, other reductions in the branchial sac and
body musculature in the latter genus are greater than in Adagnesia antarctica
which retains the biramous branchial papillae of Caenagnesia and may be
considered phylogenetically intermediate between Caenagnesia and Agnesia.
ADAGNESIA oOPAcA Kott
Adagnesia opaca Kott, 1963, p. 76.
Specimens examined.—Queensland Museum (Reg. No. G4907): coll. W.
Stephenson et al., Moreton Bay, Queensland (numerous. specimens).
Australian Museum: coll. J. MacIntyre, 16.6.65, 140 m, off Cronulla, N.S.W.
(fragments only).
Remarks.—This is a particularly large species (3 to 4 cm diameter) and
represents the most highly specialized genus of the family in regard to its
closing mechanism. The test is thin and completely rigid and brittle with
sand. On both sides of the apertures folds of test form lips so shaped
that the excurrent aperture is directed upwards and the incurrent aperture
is directed downwards toward the substrate on which the animal lies. The
body musculature is correspondingly specialized and bears a relationship to
that of Agnesia glaciata. Only a limited number of circular muscle bands
are present around the siphons, superficial to short radiating longitudinal
bands which extend no further than the base of the siphons. Both anterior
and posterior to the circular muscles of both siphons, there are very strong
transverse bands extending across the dorsal line from the base of the
protective folds of test. Their contraction draws these folds together. The
general body musculature is reduced to very short muscle bands around
the anterior, dorsal, posterior and ventral borders of both sides of the
body. These short bands along the dorso-lateral border are arranged in
parallel to the long axis of the body and may represent the remnants of the
distal portions of longitudinal bands which radiate from the siphons. The
remaining short muscle bands which appear to be deeper in the body wall
than those along the dorsal border, are probably homologues of similar
muscles in Agnesia glaciata and represent the remnants of the inner circular
body musculature. About 30 transverse vessels, each supporting about 32
bifid papillae, alternate with single rows of about 60 infundibula. However,
although there are triangular languets on each transverse vessel to the left
of the dorsal line large languets alternate with small languets suggesting
that those transverse vessels bearing the latter have developed later as a
PATRICIA KOTT a5)
result of proliferation of the branchial sac. In the posterior part of the
branchial sac this proliferation can be observed occurring and_ single
infundibula tend to subdivide into two to increase the number of spirals
in each row and then further subdivision occurs to increase the number
of rows of infundibula so that 4 spirals, arranged in a square, result from
the subdivision of a single spiral infundibulum. In association with the
highly developed closing mechanism the gonoducts are directed anteriorly
and the species is probably viviparous.
Occurrence.—Moreton Bay, Queensland (Kott 1963); off Cronulla.
N.S.W., 140 m (New Record).
Discussion.—The reduction of body musculature has proceeded beyond the
condition found in Adagnesia antarctica and indeed, beyond the degree of
reduction and specialization of this musculature known in Agnesia spp.
However the condition of the dorsal lamina and branchial papillae, and
the numbers of branchial papillae in relation to the numbers of infundibula
in each row establish its relationship to Adagnesia antarctica from which
it has diverged by modifications of the musculature to operate a highly
specialized closing mechanism, by the secondary development of numerous
transverse vessels in the branchial sac and proliferation of the numbers of
infundibula in each row and by a corresponding increase in the numbers
of branchial papillae on the transverse vessels. The similarity of the branchial
sac of this species to that of Caenagnesia spp. is due to this proliferation of
infundibula’ and transverse vessels together with the retention of biramous
branchial papillae. However the reduced number of branchial papillae
associated with each infundibulum in the present species together with the
condition of the dorsal lamina clearly distinguish the species from
Caenagnesia. The large numbers of infundibula in the branchial sac is a
secondary development perhaps related to the large size characteristic of
individuals of this species.
ZOOGHOGRAPHY
The widely dispersed records of Agnesia glaciata suggest that this
represents an ancient species with a well established cosmopolitan distribu-
tion, and that today records of this species from the Pacific, Antarctic and
Indian Oceans may represent relict populations now isolated. The occurrence
of 2 species of the family exclusively in the Antarctic also probably represent
relict populations now isolated by the submergence of land or submarine
bridges. Despite the fact that it is the phylogenetically primitive genus
Caenagnesia which is today endemic in the Antarctic it is not thought
possible for the family to have developed there and to have then radiated
northwards as far as northern boreal waters. If this had occurred one would
not expect, in isolated areas, so very few species. It is the wide distribution
and morphological homogeneity of Agnesia glaciata which suggests that
the species (and the family) was well established a long time ago and
persists today as a highly successful relict form.
References
ARNBACK CHRISTIE-LINDE, AUuGusTA, 1938.—Ascidiacea. Further zool. results Swed.
Antarct. Exped. 1901-03, 3(4): 1-54.
HuntTsMAn, A. G., 1912.—Ascidians from the coasts of Canada. Trans. r. Can. Inst.,
9: 111-148.
, 1912a—Holosomatous ascidians from the coast of western Canada. Contr.
Can. Biol. Fish. 1906-1910 : 103-185.
—_
456 A REVIEW OF THE FAMILY AGNESIIDAE HUNTSMAN
Kort, Patricia, 1952.—Ascidians of Australia. 1. Stolidobranchiata and Phlebobranchiata.
Aust. J. mar. freshwat. Reées., 3 (3): 206-333.
, 1954.—Tunicata. Rep. Series B, B.A.N.Z. antarct. Res. Hxped. 1929-1931,
1(4) : 121-185.
, 1963.—Adagnesia opaca gen. nov., sp. nov. a remarkable ascidian of the
family Agnesiidae, from Moreton Bay, Queensland. Pap. Dep. Zool. Univ. Qld.,
(3) 8 TEH08e
, 1969.—Antarctic Ascidiacea; a monographic account of known species with
additions based on collections made under U.S. Government auspices 1947-1965.
Antarctic Research Series, 138: 1-220. :
MICHAELSEN, W., 1898.—Vorlaufige Mitteilung tiber einige Tunicaten aus dem
magalhaenischen Gebiet sowie von Siid-Georgien. Zool. Anzg., 21: 363-871.
, 1900.—Die Holosomen Ascidien des magalhaenisch—stidgeorgischen Gebietes.
Zoologica. Stuttg., 12(31) : 1-148.
, 1907.—Tunicaten. Ergebnisse der Hamburger Magalhaenischen Sammelreise
1892-1893, Lfg. 8, No. 5, Hamburg (L Friederichsen & Co): 1-84.
, 1912.—Die Tethyiden (Styeliden) des Naturhistorischen Museums zu Hamburg,
nebst Nachtrag und Anhang einige anderen Familien betreffend. Jb. Hamb. wiss.
anst., 28(2): 109-186.
Mitxiar, R. H., 1955—On a collection of Ascidians from South Africa. Proc. zool. Soc.
Lond., 125(1): 169-221.
, 1955a.—Ascidiacea. Rep. Swed. deep sea exped. Zoology 2(18): 223-236.
, 1960.—Ascidiacea, “Discovery” Rep. 30: 1-160.
, 1962—Further descriptions of South African Ascidians. Ann. 8S. Afr. Mus.,
46(7): 113-221.
Oxa, A., 1915—Hine neue Ascidienart aus der Gattung Agnesia Michaelsen. Annotnes
zool. jap., 9: 1-6.
, 1929.—Hine zweite japenische Art der Gattung Agnesia. Proc. imp. Acad.
Japan, 5: 152-154.
Ritter, W. E., 1913.—The simple ascidians from the north-eastern Pacific in the
collection of the United States National Museum. Proc. U.S. natn. Mus., 45: 427-505.
Van Name, W. G., 1945.—The North and South American Ascidians. Bull. Am. Mus.
nat. Hist., 84: 1-476.
TYPE SPECIMENS IN THE MACLEAY MUSEUM
UNIVERSITY OF SYDNEY
III. Brrps
P. J. STANBURY
School of Biological Sciences, University of Sydney
[Read 27th November, 1968]
INTRODUCTION
There are about 9000 bird skins in the Macleay Museum at the University
of Sydney. At least 57 of these are types or belong to a type series. This
paper lists them together with associated pertinent information. As noted
in the other papers in this series (Stanbury 1968, Goldman, Hill and Stanbury
1969) further types may lie unrecognized in the Museum.
Tun Birp Types
Some of the bird types were collected on the “Chevert” expedition to
north Queensland and New Guinea waters in 1875 (see Macmillan 1957).
Others were obtained from collectors in the northern parts of Australia. The
new species. were described by G. Masters (the first curator of the Macleay
Museum*) and by E. P. Ramsay.
A few of the type specimens are mounted but most are stuffed skins.
All are kept in the dark in air-tight drawers.
In the table that follows the BT numbers refer to the numbers of the
compartments in which the specimens are stored. At present there is no
complete register of birds; a register has been started but it is unlikely
to be completed before the end of 1969. When the register is complete
each type specimen will be numbered with the prefix B. (The BT number
will appear in brackets on the same label.)
The classification in the table is that devised by A. Wetmore of the
Smithsonian Institute, Washington, in 1960.
The reference to the original description of each type specimen is given.
In addition the revised name used by G. M. Mathews (in the two parts of
Systema Avium Australasianarum, published in 1927 and 1930 by the British
Ornitholigists Union) is given. Reference to the page quoted will usually
provide a list of earlier synonymous names.
The Australian Museum holds some types of which the Macleay Museum
also holds individuals of the same species collected from the same locality
in about the same year.
In view of the relations of Sir William Macleay with the Australian
Museum and with the University of Sydney it is possible that some of such
specimens in the Macleay Museum are part of a type series. I have therefore
labelled these “type series?”. In these instances the reference number of
the Australian Museum type specimen also is given in the table.
1The curators of the Macleay Museum were G. Masters 1888-1912; J. Shewan
1913-1933; K. HE. W. Salter 1934-1944; J. R. Henry 1945-1958; Elizabeth Hahn 1958-1962;
and Jennifer M. EH. Anderson 1963-1966; all of whom have contributed either directly
or indirectly to this list. I am especially grateful to my immediate predecessor, Mrs.
J. Anderson, for her work in this regard.
PROCEEDINGS OF THE LINNEAN SociETy oF New SoutH WALES, VoL. 93, Part 3
III BIRDS
TYPH SPECIMENS IN THE MACLBAY MUSEUM.
458
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TYPE SPECIMENS IN THE MACLEAY MUSEUM. III BIRDS
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P. J. STANBURY 461
Acknowledgements
I wish to express my thanks to the Curator of Birds at the Australian
Museum Mr. H. J. de S. Disney, and especially to Mr. K. A. Hindwood,
Honorary Associate of the same Museum, who very kindly sorted through a
selection of suspected type specimens.
IT am also grateful to Mrs. Karen Paynter and to Miss Kerry Smith for
their proficient technical assistance in the compiling of the several drafts
of this list.
References
GotpMAN, Jupy, Him, L. and SrAansury, P. J., 1969.—Type specimens in the Macleay
Museum, University of Sydney. II. Amphibians and reptiles. Proc. Linn. Soo. N.S.W.,
93 : 427-438.
MAomILLAN, D. N. S., 1957.—“‘A Squatter Went to Sea.” Currawong Publishing Company,
Sydney.
Maruews, G. M., 1927, 1930.—“Systema Avium Australasianarum.” Parts I and II,
1047 pp. British Ornithologists Union.
Sransoury, P. J., 1968—Type specimens in the Macleay Museum, University of Sydney.
I. Fishes. Proc. Linn Soc. .N.S.W., 93: 201-210.
a
TYPE SPECIMENS IN THE MACLEAY MUSEUM
UNIVERSITY OF SYDNEY
IV. MAMMALS
P. J. STANBURY
School of Biological Sciences, University of Sydney
[Read 27th November, 1968]
INTRODUCTION
The Macleay Museum has nearly 1,200 mammalian specimens. At least
8 of these are types. As noted in the earlier papers in this series (Stanbury
1968, Goldman, Hill and Stanbury 1969, Stanbury 1969) further types may
lie unrecognized in the Museum.
Top MAMMALIAN TYPES
Six of the mammalian types were collected on the “Chevert” expedition
to New Guinea in 1875 (see Macmillan 1957). The other two, from North
Western Australia, were collected by W. W. Froggatt, one of Sir William
Macleay’s collectors.
All but one of the specimens are mounted or stuffed skins. Sminthopsis
froggatti is stored in spirit. AJl the specimens are stored in the dark. Some
fading is apparent on most of the specimens. One, Dorcopsis beccarii has lost
about 40% of its fur.
RARE MARSUPIALS
The Macleay Museum also possesses a number of rare marsupial specimens
which are not types but which now are extinct or virtually so: for example
Onychogalea fraenata (Lower Murrumbidgee, N.S.W.), Myrmecobius rufus
(S. Australia), Antechinus apicalis (King Georges Sound, W. A.). and
Thylacinus cynocephalus (Tasmania).
Acknowledgements
I am indebted to the previous curators of the Museum, and especially
to Mrs. J. Anderson (1963-1966). I wish also to thank Drs. W. D. L. Ride
and J. L. Bannister of the Western Australian Museum who checked the first
draft of this list and made many helpful suggestions.
References
GoLDMAN, JuDy, Him, L. and Stransury, P. J., 1969—Type specimens in the Macleay
Museum, University of Sydney. II. Amphibians and reptiles. Proc. Linn. Soc. N.S.W.,
93 : 427-438.
MAocmit3tan, D. N. S., 1957.—“A Squatter went to Sea.” Currawong Publishing Company,
Sydney.
Stanpury, P. J., 1968—Type specimens in the Macleay Museum, University of Sydney.
I. Fishes. Proc. Linn. Soc. N.S.W., 93: 201-210.
STANBuRY, P. J., 1969—Type specimens in the Macleay Museum, University of Sydney
III. Birds. Proc. Linn. Soc. N.S.W., 93 : 457-461.
PROCEEDINGS OF THE LINNEAN Society oF NEw SouTH WALES, VoL. 93, Part 3
463
P. J. STANBURY
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HATCHING AND LARVAL DEVELOPMENT OF HAPLOSTOMELLA
AUSTRALIENSIS GOTTO (COPEPODA, FAM. ASCIDICOLIDAE),
A PARASITE OF THE ASCIDIAN STYELA ETHERIDGII HERDMAN
D. T. ANppRSoN and G. T. Rossrrer
School of Biological Sciences, Unwersity of Sydney
[Read 27th November, 1968]
Synopsis
H. australiensis produces paired cylindrical ovisacs each containing 70-100 eggs.
Hatching occurs as a lecithotrophic, positively phototropic nauplius which swims
briefly then moults to a demersal first copepodite. This stage exhibits sexual dimor-
phism and is probably the infective stage of the life cycle. Hatching is osmotic in
the usual copepod manner and is stimulated by light and mechanical agitation. The
obligatory light stimulus ensures that hatching occurs after the ovisac is discharged
from the host. The response to mechanical agitation ensures a synchronous hatching
which ruptures the tough wall of the ovisac. The hatching mechanism is specialized
and the larval development is abbreviated compared with those of other ascidicolous
copepods.
INTRODUCTION
During the course of a general survey of copepods of the Sydney area, in
search of species suitable for embryological study, the authors collected
specimens of a large worm-like parasitic copepod inhabiting the oesophagus
and stomach of the simple ascidian Styela etheridgii Herdman. The parasite
was identified by Dr. R. V. Gotto of Queen’s University, Belfast, as a new
species of the genus Haplostomella Chatton and Harant 1924, family
Ascidicolidae. The four previously described species of Haplostomella have
all been collected from northern hemisphere localities, where they occur in
compound ascidians. The new species, named H. australiensis Gotto, has been
described by Gotto (1968). In the present paper, we present observations on
the general breeding activity, hatching and larval development of this animal.
MATERIALS AND METHODS
Adult females of H. australiensis were collected at intervals during June,
July and August, 1967. The animals were taken from specimens of Styela
etheridgvi living at low tide level on Bottle and Glass Rocks on the south
shore of Sydney Harbour. Records were kept of the frequency of occurrence
of egg masses attached to the animals, and the later embryonic development,
hatching and first two larval stages were studied using egg masses cultured
in the laboratory.
The egg masses were cultured in glass tubes, the lower ends of which
were closed with fine nylon gauze and immersed in well-aerated, filtered
sea water. The water was changed regularly and tubes cleaned of algal
film every 2 days. The cultures were maintained at 20°C. The water temper-
ature in the natural habitat of the copepods at the time of collection ranged
from 13 to 18°C.
_ The aquarium containing the culture tubes was at no time subjected to
direct sunlight, but the laboratory was illuminated through windows on its
northerly aspect during the day. Fluorescent lighting supplemented natural
PROCEEDINGS OF THE LINNEAN Society or NEw SouTH WALES, Vot. 93, Part 3
D. T. ANDERSON AND G. T. ROSSITER 465
Ulumination during working hours. At all times, the culture tubes were
shaded from light from above by a sheet of white paper placed over their
upper ends. Mention is made of these facts because it was subsequently
determined that the embryos are sensitive to bright light stimulation as a
causative factor in hatching.
The sequence of events in hatching was observed for individual embryos
and for whole egg masses. The finer details of hatching of individual embryos,
including the rupture of egg membranes and accurate measurements of
changes in volume, were recorded by mounting eggs in sea water on a cavity
slide sealed with a cover slip and observing hatching by phase contrast
microscopy.
The sensitivity of the embryos to bright light stimulation was studied
using a sharply focussed microscope lamp to deliver either flashes or longer
periods of bright light stimulation. By this technique, the days on which
embryos could be induced to hatch by stimulation with bright light were
determined, relative to the day of hatching of unstimulated embryos of the
Same age. This difference is referred to in the following account as days
before “normal” hatching. —
The hatching response of embryos in solutions of different osmotic
strengths was also studied, in accordance with the view expressed by Marshall
and Orr (1954) and Davis (1959) that a common osmotic mechanism
underlies hatching in copepods. Embryos were immersed in hypotonic or
hypertonic sea water in Petri dishes brightly illuminated under a binocular
100
Zadult female
parasites
carrying egg
masses
/ascidians
containing an
adult female
parasite
50 50
8/6 22/6 246 8/6 22/6 24/6
B
Fig. 1 A, % occurrence of adult female Haplostomella in three samples of Styela from
a single population; B, % of parasites in each sample bearing egg masses.
microscope. Newly emerged nauplii, or the embryos if hatching failed to
occur within a fixed period, were then transferred back to normal sea water.
Controls remained in normal sea water throughout. Hypotonic sea water, -
osmotically equivalent to a 2-4% salt solution, was prepared by diluting sea.
water (3:5% salt solution) to 70% with distilled water. Hypertonic sea
water (75% salt solution) was prepared by adding 4 gm. sodium chloride
per 100 ml. to normal sea water.
RESULTS
Occurrence and breeding of adult females
The percentage occurrence of H. ausiraliensis females in the ascidian
Styela etheridgii in three separate samples collected at Bottle and Glass
466 DEVELOPMENT OF HAPLOSTOMELLA AUSTRALIENSIS GOTTO
Rocks is shown in Fig. 1A. The percentage, in each sample, of females
which carried egg masses is Shown in Fig. 1B. The decline in the percentage
of ovigerous females in the second and third samples suggests a trend towards
termination of a breeding cycle. The number of ascidians examined in the
samples were 16, 75 and 88 respectively.
In none of the ascidians examined was there more than one adult female
Haplostomella. The minimum length of the specimens was 4 mm., while the
average length was 5 mm.
Culturing of egg masses and larvae
Most of the difficulties previously associated with the laboratory culture
of egg masses from parasitic copepods (Wilson 1905, 1907a, 1907b, 1911a,
19116) were experienced again in the present work. From the culture of
32 egg masses, only 6 pairs of masses already containing embryos at an
advanced stage of development when collected remained viable until hatching.
From 38 of these masses, embryos which had received no form of artificial
stimulation hatched after 9, 18 and 17 days of culture respectively. Egg
masses with embryos at very early stages of development, for example,
cleavage or gastrulation, remained viable in culture for about 15 days.
Maintenance of the hatched larval stages in the laboratory also proved
difficult. About 95% of the nauplii hatched from egg masses in the laboratory
failed to survive the first moult into a first copepodite stage. Because of the
low survival rate, the few first copepodites which did emerge were fixed
while intact, and development was not followed beyond this stage.
The egg masses
H. australiensis, like many copepods, lays its eggs into paired cylindrical
ovisacs attached to the genital apertures of the female. The eggs when laid
are spherical and 180u in diameter. Seventy to 100 eggs are closely packed
within each ovisac, which measures about 3 mm. x 0-8 mm. The eggs tend to be
flattened slightly in the early stages of development, due to their close packing.
They then gradually assume a regular ellipsoidal shape, 200 long and 160p
in diameter.
The ovisacs function as brood chambers and remain attached to the
female at least until the embryos reach a late stage of development. The
period of development is estimated to be 6-8 weeks, taking into account the
observation of 2 weeks of early development and 24 weeks of later development
of embryos in culture. No ovisacs with embryos at an intermediate stage of
development were collected.
The newly hatched larva
The newly hatched nauplius of H. australiensis, 200u in length, is
illustrated in Fig. 2G. The nauplius is free-swimming and lecithotrophic. The
antennules, antennae and mandibles all bear swimming setae, but feeding
spines are absent. The labrum is also poorly defined. The post-naupliar region
is large and swollen, but shows no external signs of segmentation or limb
rudiments. The nauplius is markedly phototropic and is a very rapid
swimmer.
The sequence of events in hatching
Towards the end of embryonic development, for almost a week before
hatching, the embryo shows peristaltic movements of the gut and irregular,
rather infrequent twitching of the body. Just before hatching is initiated,
D. ‘I’. ANDERSON AND G. T. ROSSITER 467
there is a marked increase in activity. For a period of one or two minutes,
the embryo exhibits vigorous movements of the whole body and especially
of the three pairs of limbs.
caudal spine
yolky midgut nauplius eye
inner membrane
antennule
antenna
mandible
Fig. 2. The hatching sequence of H. australiensis (for description, see text).
468 DEVELOPMENT OF HAPLOSTOMELLA AUSTRALIENSIS GOTTO
After this phase of vigorous movement, the nauplius becomes quiescent
and a thin film of fluid appears between the nauplius and the egg membranes
(Figs 2A, 2B). The increasing volume of the fluid causes the membranes to
swell, usually in the equitorial plane. The outer egg membrane ruptures,
revealing the transparent inner membrane (Figs 2C, 2D). During this
phase of swelling, the naupliar limbs move passively with the expanding inner
membrane until they are in the extended swimming position (Fig. 2E).
The volume of fluid within the inner membrane continues to increase until
the membrane forms a sphere 270» in diameter (Fig. 2F). The average time
for expansion is 1-2 minutes; and the volume increases by a factor of 4:°3=1.
The nauplius, which fills the egg membranes before swelling begins, shows
no increase in volume during the hatching process.
The nauplius remains quiescent within the inner membrane for a further
period of 1-2 minutes after the inner membrane has ceased to expand. The
inner membrane is then ruptured as the nauplius swims rapidly forwards
(Fig. 2G). The nauplius, freed from its egg membranes, swims immediately
to the surface and towards the light.
Physical factors influencing hatching
(i) Exposure to bright light:
The marked phototropism of the newly hatched nauplius is preceded
by sensitivity to bright light for a number of days before hatching. During
those days, muscular twitching of the embryo can be evoked on exposure
to flashes of bright light, and the embryo can be stimulated to hatch pre-
maturely by continuous exposure to bright light. The sensitivity to light
of embryos from a single egg sac, measured for five days preceding normal
hatching, is summarized in Table I.
Tasre I
Sensitivity to light of embryos from a single ovisac of Haplostomella australiensis
Days Number Period of exposure to _ Average -
before of bright light required exposure
normal embryos to initiate hatching period
hatching tested (minutes) (minutes)
5 : 3 One hatched after 180 > 180
minutes. Two failed to
hatch in this time
4 3 120-180 150
3 10 30-90 80
2 gs 4-20 15
1 25 1-7 3
As can be seen, the exposure time required to induce hatching falls
rapidly as the day of normal hatching approaches. The sensitivity of the
twitch response to a flash of bright light also increases during the final days
of embryonic development.
(ii) Mechanical agitation:
Mechanical agitation also hastens the onset of hatching. In one embryo
two days before normal hatching, mechanical agitation with needles under
bright light caused hatching to commence after 3 minutes. The average time
D, T. ANDERSON AND G. T. ROSSITER 469
of exposure to bright light required to initiate hatching in embryos of this
age is 15 minutes, and the minimum time is 4 minutes (Table T). A second
embryo, one day before normal hatching, began to hatch while being removed
from the ovisac with needles, even before being exposed to bright light.
For 5 embryos spread out in a Petri dish and exposed to bright light,
15 minutes elapsed between the first sign of hatching in the first embryo
to hatch and in the last to hatch. For 20 embryos from the same ovisac,
kept in contact inside part of the ovisac, the interval was only 6 minutes.
In this case, the vigorous prehatching movements of the first embryo to
hatch caused mechanical agitation of adjacent embryos and a rapid spread
of the hatching response ensued.
The synchronization of hatching
Hatching under bright light stimulation was induced and observed for
three intact ovisacs. In each case, the sequence of hatching and its timing
for individual embryos within the ovisac was found to be the same as that
previously described for individual embryos removed from their ovisacs and
studied in isolation. Moreover, the functional significance of the sensitivity of
the hatching response to mechanical agitation was revealed by these observa-
tions. The vigorous prehatching movements induced by the bright light
stimulus began at first in only a few scattered embryos, but spread quickly
through adjacent embryos until the whole ovisac vibrated. This mass activity
persisted for about 3 minutes, and then almost all of the embryos simul-
taneously became quiescent as the 1-2 minute phase of expansion of the egg
membranes began. The simultaneous increase in volume of the inner
membranes, in addition to rupturing the outer membranes, generated a
collective pressure sufficient to rupture the ovisac longitudinally. Most of
the embryos, still quiescent within their expanding inner membranes, spilled
out through the rupture and completed their hatching outside the ovisac.
Those remaining within the ovisac hatched and escaped through the longi-
tudinal slit.
The importance of synchronous hatching in promoting escape from the
ovisac-was confirmed by another experimental observation. Five embryos, one
day before normal hatching, were loosely packed and sealed off in part of an
ovisac. Simultaneous hatching was then induced by exposure to bright light,
but the expansion of the egg membranes was able to be accommodated without
rupture of the surrounding ovisac wall. The nauplii thus escaped within the
closed ovisac. Although they made vigorous swimming movements, the nauplii
were unable to rupture the tough ovisac wall and escape into the surrounding
water.
Natural hatching
The hatching of embryos in the laboratory without bright light stimula-
tion was not observed directly, but several batches of nauplii hatched in
culture from ovisacs which had been observed to be intact on the previous day.
Such hatching was noticed at about 9 a.m., by which time most of the nauplii
had entered the quiescent period preceding the moult to a first copepodite
(see below). This quiescence follows a 2-3 hour period of activity, indicating
that the nauplii had hatched synchronously at about dawn. The associated
ruptured ovisacs gave further indication that natural hatching proceeds
in the same synchronous manner as experimentally induced hatching.
470 DEVELOPMENT OF HAPLOSTOMBLLA AUSTRALIENSIS GOTTO
Hatching in hypotonic and hypertonic sea water
(i) Hypotonic sea water:
The hatching sequence was not altered by immersion of eggs in hypotonic
sea water, but the rate of hatching was accelerated (Fig. 3). The duration
of both the phase of prehatching activity and the phase of expansion of
| solution time of exposure to bright light (minutes)
te) 10 20 30 40 50 60
normal
seawater
petals
(3-5 /saline)
hypotonic
seawater
(2-4/saline)
hypertonic
seawater
(7-5/saline)
Fig. 3. The timing of light-stimulated hatching of H. australiensis in hypotonic and
hypertonic sea water, for embryos 2 days before normal hatching. Hach shaded block
represents the time for one embryo to complete the hatching sequence.
the inner membrane were shortened. When returned to normal sea water
after hatching, the nauplii which had been hatched in hypotonic sea water
did not differ in behaviour or further development from those hatched in
normal sea water.
(ii) Hypertonic sea water:
Following immersion in hypertonic sea water, both the hatching sequence
and the rate of hatching were altered. In response to the stimulus of bright
light, the embryos began their prehatching movements in the usual way, but
the swelling of the egg membranes was completely inhibited. Only one embryo
out of nine attained hatching in the hypertonic solution, in approximately
normal time (Fig. 3). Of the remaining 8, three underwent delayed hatching
after return to normal sea water, while the remainder failed to hatch.
Although prehatching activity was prolonged, no swelling of the egg
membranes ensued even after return to normal sea water.
Larval development
(i) The nauplius:
The nauplius, described and illustrated above, remains active at the
surface of the water for 2-3 hours after hatching. It then sinks and becomes
quiescent, entering the moult to the first copepodite stage.
D. T. ANDERSON AND G. T. ROSSITOR 47]
welll tm, antennule
a antenna
mandible
XA -- maxilliped
second thoracic
limb
‘third thoracic
limb
fourth thoracic
limb
caudal fork
1009p
antennule
antenna
mandible
labrum
maxilliped
second thoracic
limb
third thoracic
limb
fourth thoracic
lamb
caudal fork
Fig. 4. The first copepodite of H. australiensis. A, female, ventral view: B, female,
_ lateral view; C, male, ventral view; D, male, lateral view.
472 DEVELOPMENT OF HAPLOSTOMELLA AUSTRALIENSIS GOTTO
The nauplius swims in the manner typical of newly hatched copepod
nauplii (Gauld, 1959), by rapid, synchronous beating of the three pairs of
naupliar limbs. Swimming follows a spiral course. At no time does the
nauplius display the slower, rhythmic movements typical of the later naupliar
stages of planktonic copepods. The main propulsion comes from the action
of the long, setose antennae and mandibles, with a lesser contribution from
the shorter antennules.
(ii) The first copepodite:
Three to four hours after the onset of quiescence, the nauplius moults,
yielding a first copepodite stage (Fig. 4). This larva is 280-300» long. The
naupliar appendages are greatly reduced, while the first three pairs of bira-
mous trunk limbs are exposed as functional swimming limbs. The copepodite
is not phototropic, and swims at or near the bottom of its container. Probably
this stage or the next is the stage which enters the ascidian host.
The first copepodite of H. australiensis already shows sexual dimorphism.
The male (Figs 4C, 4D) can be diagnostically distinguished from the female
(Figs 4A, 4B) by the antennules, which are distinct and segmented in the
male, small and inconspicuous in the female, a difference reflecting the
specialized later function of the antennules as claspers in male ascidicolid
copepods (Lang, 1948). In addition, the remaining naupliar limbs and trunk
limbs of the male are generally larger and more setose than those of the
female, and a greater space is observed between the maxillipeds and first pair
of swimming limbs.
Neither the copepodite stage nor the adult female show traces of
maxillules or maxillae. In view of the vermiform character and reduced
limbs of the adult female, it is clear that further female larval stages would
show the acquisition of two more pairs of trunk limbs, followed by reduction
and simplification of the five pairs. The adult male of H. australiensis has
not been identified, but probably resembles the Cyclops-like male of Ascidicola
(Gotto, 1957) and has a more typical later larval development.
DISCUSSION
Duration of development
The apparent duration of embryonic development in H. australiensis,
6-8 weeks, is comparable with those reported by Wilson (1905, 1907a, 19076,
1911a, 19116) for a great number of parasitic copepods. Such periods are
long compared with the few days of embryonic development in Cyclops and
the even shorter 1—2 days in Calanus (Marshall and Orr, 1954). The extended
embryonic development of parasitic copepods is a reflection of a combination
of larger eggs and prolonged direct development to a late larval stage before
hatching. Ascidicola rosea, with a smaller egg, takes 18 days (Gotto, 1957),
and Gonophysema 19-20 days (Bresciani and Litzen, 1961).
The hatching mechanism
The sequence of events in hatching, and the operation of the osmotic
hatching mechanism in H. australiensis are comparable with those described
for marine planktonic copepods by Marshall and Orr (1954) and for fresh
water copepods by Davis (1959). Each of these shows the phase of vigorous
activity just before hatching, the osmotic expansion of the inner membrane
and rupture of the outer membrane, and the quiescence of the embryo during
expansion of the membranes.
D, T. ANDERSON AND G. T. ROSSITER 473
The observation for H. australiensis that immersion in hypotonic sea
water affects the rate of hatching but not the time of onset of hatching,
suggests that the hatching mechanism can be resolved into two components.
The first is a light-sensitive component which, when activated, sets in motion
the second, an osmotic component. It can be inferred that osmotic stress
is set up between the contents of the space between the embryo and its
membranes on the one hand, and the surrounding water on the other, as a
result of the prehatching activity of the embryo. This activity could result
either in secretion of an osmotically active substance or in a change in the
structure and permeability of the inner membrane, or both. All that is
obvious at the present time is that the activity has a light-sensitive trigger
and that the osmotic stress resulting from the activity leads to membrane
swelling and escape of the nauplius.
Hatching in natural conditions
Gotto (1957) observed through the transparent wall of Corella parallelo-
gramma (Muller) that Ascidicola rosea, which normally inhabits the
oesophagus of the host, migrates to the stomach of the host before releasing
its ripe egg masses. Here the inner membranes of the eggs swell and burst
forth from the clustered outer membranes, but remain intact around the
nauplii during passage through the intestine and rectum. Escape of the
nauplii from the swollen inner membranes occurs in the exhalent water
current of the host.
Styela etheridgii has an opaque test, and events intervening between
release of the egg sacs by Haplostomella australiensis and ensuing natural
hatching of the nauplii can only be inferred. In almost 200 ascidian hosts
examined, however, no trace was found of detached ovisacs or hatched nauplii
of the parasite. This suggests that the ovisacs released by the parasite are
carried quickly through the gut and out of the exhalent siphon of the host
before hatching takes place. Furthermore, no ovisacs were obtained that
hatched normally in less than 9 days. Presumably, ovisacs closer to hatching
than 9 days have already been released. The light-sensitivity of hatching is
probably significant in preventing hatching before the ovisac has escaped
the host since, in contrast to A. rosea, the nauplii hatch from their inner
membranes as soon as these have swollen and would be exposed to digestive
enzymes within the host. Similar light sensitivity in hatching is common
among endoparasitic trematodes and cestodes (Smyth, 1961).
Sensitivity to mechanical stimulation probably plays a different role.
When the ovisac is released from the female, it is closed off as a tough-
walled bag. Experimentation shows that rupture of this bag, essential for the
escape of the nauplii, depends on synchronous swelling of the contained egg
membranes. The prehatching activity which leads to swelling, although
dependent on light stimulation, is accelerated in its onset by mechanical
stimulation. Synchrony of the hatching process is assured by this response,
since mechanical stimulation spreads rapidly in an ovisac aS soon as one
embryo makes the prehatching response to light.
During a period of 5 days at 20°C., the embryos become more and more
sensitive to both light and mechanical stimulation, until a slight stimulus
of either kind will set off synchronous hatching. Presumably this extended
gradient of increasing sensitivity permits some margin of delay in escape
from the host, while at the same time ensuring eventual hatching even in
conditions of relatively low illumination which might be encountered.
M
474 DEVELOPMENT OF HAPLOSTOMELLA AUSTRALIENSIS GOTTO
Synchronous hatching in parasitic copepods with sac-like ovisacs may be
a common phenomenon. Wilson (1911a, 19110) noted split, empty ovisacs
of lernaeopodid and ergasilid copepods after overnight mass hatchings, and
also observed that the ovisac ruptured in Hrgasilus as the egg membranes
swelled. :
Larval stages
It is well known that the notodelphyiniform copepods inhabiting
ascidians hatch as a phototropic, free-swimming nauplius with reduced limbs
and an enlarged, ovoid, yolk-filled body (Canu, 1892; Gray, 19336; Gurney,
1933; Lang, 1948; Gotto, 1957; Gage, 1966). The Enterocolidae also have
a nauplius of this type, although lacking the positive phototropy displayed
in the notodelphyiniform species (Canu, 1892; Gray, 1933a; Lang, 1948).
The further development of the nauplius has been studied in only a few
species (Ascidicola, Notodelphys, Enterocola; Gotto, 1957; Gage, 1966; Canu,
1892) but has been consistently found to pass through four free-swimming
naupliar stages and two copepodite stages, of which the second gradually
adopts a demersal habit and is probably the infective stage. The free-
Swimming period of these species is several days.
Haplostomella australiensis has a much more abbreviated larval develop-
ment. The nauplius conforms to the usual type, being similar to that of
Ascidicola rosea, but is free-swimming for only a few hours before moulting
to the demersal first copepodite stage. Furthermore, this stage already
manifests sexual dimorphism, which probably indicates that it is the infective
stage, as in lernaeopodids (Wilson, 1911@). The only comparable. brevity
known at the present time for the free-swimming stages of a copepod
parasitizing ascidians is that of Gonophysema gullmarensis. This aberrant
species also has a single, brief naupliar stage and a single copepodite stage
infective to new hosts (Bresciani and Liitzen, 1961).
Acknowledgements
We would like to thank Miss S. A. McPhail and Miss S. A. Christmas
for technical assistance during the course of this work, Dr. R. V. Gotto,
Zoology Department, Queen’s University, Belfast, Northern Ireland, for
identification of the parasite and Miss I. Bennet, School of Biological Sciences,
University of Sydney, Miss E. Pope, Australian Museum, Sydney and Mrs.
P. Mather, Department of Zoology, University of Queensland, for identification
of the host. The work was supported by research grants from the Australian
Research Grants Committee and the University of Sydney.
References
BRESCIANI, J., and Liitzen, J., 1961—Gonophysema gullmarensis (Copepoda parasitica).
An anatomical and biological study of an endoparasite living in the ascidian
Ascidiella aspersa. II. Biology and development. Cah. Biol. mar., 2: 347-872.
CANu, E., 1892.—Les copépodes du Boulonnais. Trav. Stat. zool. Wimereux, 6: 1-345.
Davis, C. C. 1959.—Osmotic hatching in the eggs of some freshwater copepods. Biol.
Bull. Woods Hole, 116: 16-22.
GAGE, J., 1966.—Seasonal cycles of Notodelphys and Ascidicola, copepod associates with
Ascidiella (Ascidiacea). J. Zool., 150: 223-233.
GAuULD, D. T., 1959—Swimming and feeding in crustacean larvae. The nauplius larva.
Proc. zool. Soc. Lond., 132: 31-50.
GoTto, R. V., 1957.—The biology of a commensal copepod, Ascidicola rosea Thorell, in
the ascidian Corella parallelogramma (Miiller). J. mar. biol. Ass. U.K.. 36: 281-290.
, 1969.—Haplostomella australiensis, n. sp., an ascidicolous copopod from New
South Wales. Orustaceana, in press.
D. T. ANDDRSON AND G. T. ROSSITHR 475
Gray, P., 1933a.—The nauplii of Notodelphys agilis Thorell and Doropygus porcicauda
Brady. J. mar. biol. Ass. U.K., 18: 619-522.
, 19383b.—Mycophilus rosovula n. sp., a notodelphoid copepod parasitic within
B. (Botrylloides) leachii Sav., with a description of the nauplius and notes on
its habits. J. mar. biol. Ass. U.K., 18: 5623-527.
Gurney, R., 1933—Notes on some Copepoda from Plymouth. J. mar. biol. Ass. U.K.,
19: 229-804.
LAna, K., 1948.—Copepoda ‘“Notodelphyoida”’ from the Swedish west coast, with an
outline of the systematics of the copepods. Ark. Zool., 40: 1-36.
MarsHatr, S. M., and Orr, A. P., 1954—Hatching in Calanus finmarchicus and some
other copepods. J. mar. biol. Ass. U.K., 33: 393-401.
Smyru, J. D., 1962.—‘An introduction to animal parasitology.” London, English Univ.
Press.
Witson, C. B., 1905.—North American parasitic copepods belonging to the family
Caligidae. Part I. The Caliginae. Proc. U.S. nat. Mus., 28: 479-672.
, 1907a.—North American parasitic copepods belonging to the family Caligidae.
Part 2. The Trebinae and Euryphorinae. Proc. U.S. nat. Mus., 31: 669-720.
, 1907b.—A revision of the Pandarinae and Cecropinae. Proc. U.S. nat. Mus.,
31: 323-490. 7
, 1911la.—North American copepods belonging to the family Lernaeopodidae.
Proc. U.S. nat. Mus., 39: 186-226.
, 1911b.—North American copepods belonging to the family Ergasilidae. Proc.
U.S. nat. Mus., 39: 263-400.
HATCHING AND LARVAL DEVELOPMENT OF DISSONUS
NUDIVENTRIS KABATA (COPEPODA, FAM. DISSONIDAB),
A GILL PARASITE OF THE PORT JACKSON SHARK
D. T. Anpmrson and G. T. Rossrrur
School of Biological Sciences, University of Sydney
[Read 27th November, 1968]
Synopsis
D. nudiventris lays typical, uniseriate, caligid egg strings, but hatches as a
nauplius which remains attached to the egg-string by paired caudal threads and has
reduced naupliar limbs. The nauplius moults, yielding a freeswimming, demersal
copepodite which is probably the infective stage of the life cycle. The peculiarities
of hatching and development in D. nudiventris support the removal of the genus
from the Caligidae to the monogeneric family Dissonidae, proposed on morphological
grounds by Yamaguti.
INTRODUCTION
While the number of specific descriptions of copepods parasitizing fishes
in Australian waters is gradually increasing (Heegaard, 1962; Kabata, 1965,
1966), nothing has been recorded of their larval stages. The genus Dissonus,
comprising a small number of caligoid species associated with elasmobranch
fishes, is a particularly interesting one. Created by Wilson (1906), this
genus remained within the Caligidae until 1963, when it was removed by
Yamaguti (1963) to a separate family Dissonidae on the basis of a number
of small but significant morphological differences. Dissonus nudiventris was
first collected by the British, Australian and New Zealand Antarctic Expedi-
tion of 1929-1931, from Heterodontus phillipi Blainville at Hobart, Tasmania,
but the material was described and named as a new species only by Kabata
(1965). Although Kabata followed Yamaguti in assigning the species to the
family Dissonidae, the first words of his specific description read “A typical
caligid copepod—————”, emphasizing the fact that the adults of D.
nudiwentris are similar in general form to the well known caligids.
Wilson (1905, 1907a, 19076), Gurney (19384) and Heegaard (1947) have
stressed the uniformity in structure and habits of caligid larvae. The
following description of the hatching and early larval development of D.
nudiventris shows that in Yamaguti’s newly proposed family, larval develop-
ment differs markedly in the early stages from that of the Caligidae.
MATERIALS AND METHODS
Male and female adults of Dissonus nudiventris were collected from the
gill filaments of the Port Jackson shark, Heterodontus portjacksoni (Meyer).
The fishes came from the marine aquarium at Manly, N.S.W. Of four
individuals examined, one yielded 23 females and 6 male parasites, a second
65 females only and a third 1 female only, while the fourth was uninfested.
In all, fourteen egg strings of D. nudiventris containing advanced embryos
were cultured. The egg strings were found to be highly susceptible to
bacterial and protozoan attack in the conditions of the culture. Using
movements of the gut in the embryos as a convenient index of viability. it
PROCEEDINGS OF THE LINNEAN SocieTY oF NEW SOUTH WALES, VoL. 93, Part 3
D. T. ANDERSON AND G. T. ROSSITER 477
was found that the embryos remained alive for no more than five days.
Hatching was observed only if it took place during this period. Numerous
nauplii were hatched, but very few survived the first moult to the first
copepodite stage, and no larvae were carried beyond this stage.
In spite of these difficulties, which are well known for parasitic copepods
(Wilson, 1905, 1907a, 190760; Heegaard, 1947), the results obtained revealed
a unique pattern of hatching and larval development in Dissonus nudiventris,
quite unlike that of any caligid.
RESULTS
The egg string
D. nudiventris produces paired egg strings of typical caligid form. Each
ege string is cylindrical, 4-6 mm. long, 0°35 mm. in diameter, and contains
a single row of flattened, disc-shaped eggs, closely packed together with their
flattened surfaces almost touching. Each egg lies in a separate cell of the
egg string, with diameters of 350 x 320u and a thickness of 70yp. The wall
of the cell is relatively rigid and tough. As the embryos develop, it can be
seen that the flattening of the egg is a dorso-ventral one. The dorsal surface
antennule
nauplius eye
antenna
labrum
mandible
first maxilla
second maxilla
maxilliped
yolky midgut
100p
second thoracic
limb
/ third thoracic
limb
ee caudal fork
Y
yam attachment thread
Fig. 1. Dissonus nudiventris. A. The newly hatched nauplius, ventral view. B. The
first copepodite, ventral view.
of each embryo faces the ventral surface of the next embryo of the string.
Furthermore, the anterior ends of the embryos, marked in later stages by
the nauplius eyes, are also aligned. The embryos all face in one direction
along the length of the string.
478 LARVAL DEVELOPMENT OF DISSONUS NUDIVENTRIS KABATA
The newly hatched larva
The newly hatched nauplius which emerges from a cell of the egg string
in D. nudiwentris is unique among Crustacea. Although diagnostically a
nauplius, with 3 pairs of naupliar limbs and an externally unsegmented post-
naupliar region (Fig. 1A), it is non-feeding and non-swimming. The nauplius
remains permanently attached to the wall of the egg string by a pair of
fine, fibrous threads arising from the posterior end of the larva. The post-
naupliar region is elongated, and the naupliar limbs are short, devoid of
Swimming setae or feeding spines, ventrally disposed and generally similar
to the form they take in the next larval stage.
Hatching
Hatching of each individual nauplius from its cell in the egg string is
a relatively prolonged process. A variable period of activity, which may
continue sporadically for many minutes, precedes hatching. The first sign
of hatching is a slight swelling of the membrane around the embryo, accom-
panied by formation of a film of fluid between the embryo and the membrane
(Figs 2A, 2B). The embryo rounds up slightly from its previous flattened
form. Further swelling of the membrane, resisted by the tough wall of the
cell, gradually carries the nauplius out through an aperture formed in the
wall of the cell, immediately in front of the nauplius eye (Fig. 2C). The
formation of this aperture is imperceptible. Vigorous movements of the
embryo occur during this phase of hatching. The increase in volume of the
membrane is approximately fourfold.
As the nauplius is gradually extruded from its cell, the caudal threads
become apparent (Fig. 2C) and gradually extend (Fig. 2D). When the
swollen membrane breaks and the nauplius finally emerges, the threads
continue to restrain it. At this stage, they measure about twice the length
of the nauplius, which itself is about 370u long. The nauplius continues to
twitch its limbs and body after hatching, but makes no swimming movements
and shows no photosensitivity. The duration of hatching of individual nauplii
emerging from three different egg strings ranged from a few minutes to
three hours. Hatching within any one egg string was asynchronous and was
not promoted by exposure to light. It was, however, sensitive to the osmotic
concentration of the surrounding medium. Immersion of part of an egg
string, from which some nauplii had already hatched, in hypotonic sea
water (2-4% saline), caused 10 more nauplii to hatch within 5 minutes, a
degree of synchrony never observed in normal hatching. Conversely, immersion
of another part of the same egg string in hypertonic sea water (7:5% saline)
inhibited all further hatching. It can be inferred that, as in other copepods
(Marshall and Orr, 1954; Davis, 1959; Anderson and Rossiter, 1968), the
swelling of the egg membrane during hatching in D. nudiventris has an
osmotic basis.
Larval development
(i) The nauplius:
The nauplius (Fig. 1A) remains unchanged in external appearance until
it moults, after 2-5 hours, into a first copepodite. The nauplius remains
attached to the egg string by the caudal threads, and shows only sporadic
twitching. The large, curved, spatulate caudal spines of the typical caligid
free-swimming nauplius, referred to by Wilson (1905) as balancing organs,
are conspicuously absent.
A
D. T. ANDHRSON AND G. T. ROSSITER 479
(ii) The first copepodite :
The first copepodite (Fig. 2B) emerges through a longitudinal split in
the anterior dorsal midline of the naupliar cuticle, leaving the exuvia intact
and still attached to the egg string by the paired caudal threads. The first
copepodite retains the same size and general shape as the nauplius, but shows
three new features:
(a) further modification of the naupliar limbs. The antennules become
segmented and setose. The antennae become more hook-like, approximating
to their adult form. The stylet-like mandibles come into closer association
with the labrum, which itself becomes larger and pointed;
Fig. 2. The hatching sequence of Dissonus nudiventris (for description, see text).
(ob) emergence of the maxillules, as stylets associated with the labrum
and of elongated maxillae with chelate tips;
(c) emergence of three trunk segments, with a pair of long clawed
maxillipeds anteriorly on the first and two pairs of functional, biramous
swimming limbs posteriorly on the second and third segments.
The first copepodite is lecithotrophic, free-swimming and demersal,
Swimming about near the bottom of the container. There is no sign of a
frontal filament, the thread by which typical caligid copepods (the chalimus
stage) attach themselves to their hosts before completion of the larval stages
(Wilson, 1907; Gurney, 1934; Heegaard, 1947; Baer, 1952).
480 | LARVAL DEVELOPMENT OF DISSONUS NUDIVENTRIS KABATA
No sexual dimorphism is observed in the first copepodite of D. nudiventris
but, as noted by Kabata (1965), sexual dimorphism is not marked in the
adults of this species. Apart from differences in proportion in the genital
segments, the adult female differs from the male only in the absence of the
vestiges of a sixth pair of thoracic limbs.
DISCUSSION
The egg strings and hatching
The egg strings of D. nudiventris are similar to those of the many caligid
species described by Wilson (1905) and others. The sensitivity of the embryos
and larvae in conditions of artificial culture also reiterates the findings of
Wilson and of Heegaard (1947) for the eggs of many copepods parasitizing
fish. Generally, these parasites experience very efficient ventilation, the
absence of which may be a critical factor in culture.
Associated with this difficulty, the long duration of hatching from the
egg string in D. nudiventris may not be a true indication of hatching in
natural conditions. Wilson (1905, 1907) observed that hatching from a
typical caligid egg string takes place more or less simultaneously, and in
view of the synchrony of development of the eggs in the egg string of D.
nudiventris, synchronous hatching is to be expected. Since each embryo
emerges independently from its own cell in the egg string, however, the role
of synchronous hatching in rupturing the ovisac in the ascidicolid copepod
Haplostomella australiensis and other species with sac-like ovisacs (Anderson
and Rossiter, 1968) is irrelevant in D. nudiventris.
Larval development
While passage through most or all of naupliar development within the
egg membranes is a feature of lernaeid and lernaeopodid copepods parasitizing
fishes (Wilson, 191la; Baer, 1952) and of Gonophysema gullmarensis and
Haplostomella australiensis parasitizing ascidians (Bresciani and Liitzen,
1961; Anderson and Rossiter, 1968) there has been no previous record of a
non-Sswimming nauplius which hatches and yet remains attached by paired
threads to the egg string, as in D. nudiventris. This nauplius is a remarkable
exception to the rule noted by C. B. Wilson (1911@) that “all copepod
nauplii, as well those of parasitic forms as those of free swimmers, seek the
surface of the water and there swim about freely”.
The brief attached naupliar stage of D. nudiventris can be interpreted
as an exotic means of shortening the duration of free larval stages, alternative
to the prolonged direct development within the egg seen in lernaeopodids and
many other Crustacea. It contrasts strongly with the typical succession of
two phototropic, free-swimming naupliar stages and two demersal copepodite
stages seen in caligid development (Heegaard, 1947). The caligid larvae
provide a prolonged dispersal and host-seeking phase in the life cycle. The
elimination of the free-swimming naupliar stages in D. nudiventris, with
retention of the demersal first copepodite, greatly restricts dispersal,
apparently in favour of protected, more direct development. This loss may
not be important, however, since both male and female adults of D.
nudiventris can leave the host and swim freely, at least for short periods.
Associated with the loss of naupliar swimming, the naupliar limbs develop
directly towards their later adult form, and the first copepodite already
has mouthparts approximating in structure to those of the adult. Since
the larva is also lecithotrophic, slow-swimming and demersal, it seems likely
to be the immediate host-seeking and attachment stage. It follows that the
D. T. ANDERSON AND G. T. ROSSITER 48]
peculiar attached nauplius is in all probability an adaptation to completion
of the life cycle of D. nudiwentris on a single host shark, with the emergent
first copepodite stage attaching directly to the gills on to which it emerges.
This might well be advantageous in view of the solitary habits and scattered
distribution of the Port Jackson shark, leaving host transfer to the adult
during moments of host contact, e.g. in mating. Infestation by one ovigerous
female would then be sufficient to ensure rapid parasitization of a previously
unparasitized host. It is notable that of the sharks examined, one was totally
uninfested.
Acknowledgements
We would like to thank Miss 8. A. McPhail and Miss S. A. Christmas
for technical assistance during the course of this work, Professor G. H.
Satchell, School of Biological Sciences, University of Sydney, for making
available the Port Jackson Sharks, and Mr. G. C. Hewitt, Zoology Department,
Victoria University of Wellington, New Zealand, for the identification of
the parasite. The work was supported by research grants from the Australian
Research Grants Committee and the University of Sydney.
References
Anperson, D. T., and Rossiter, G. T., 1968——Hatching and larval development of
Haplostomella australiensis Gotto (Copepoda, Fam. Ascidicolidae), a parasite of
the ascidian Styela etheridgii Herdman. Proc. Linn. Soc. N.S.W., 93 : 464-475.
Barr, J. G., 1952.—‘Heology of animal parasites.” Urbana. Univ. of Illinois Press.
BRESCIANI, J., and Liitzen, J., 1961—Gonophysema gullmarensis (Copepoda parasitica).
An anatomical and biological study of an endoparasite living in the ascidian
Ascidiella aspersa. II. Biology and development. Cah. Biol. mar., 2: 347-372.
Davis, C. C., 1959.—Osmotic hatching in the eggs of some freshwater copepods. Biol.
bull. Woods Hole, 116: 15-29.
Gurney, R., 1934——The development of certain parasitic Copepoda of the families
Caligidae and Clavellidae. Proc. zool. Soc. Lond., 1934, 177-217.
HeEEGAARD, P., 1947——Contributions to the phylogeny of the arthropods. Copepoda.
Spolia Zool. Mus. Hauniensis, 8: 1-236.
, 1962.—Parasitic Copepoda from Australian fishes. Rec. Aust. Mus., 25: 149-234.
Kapara, Z., 1965.—Parasitic Copepoda of fishes. B.A.N.Z. Antarctic Research Expdn.,
1929-31. Reports, Ser. B (Zool. and Bot.), 8 (6): 1-16.
, 1966.—Copepoda parasitic on Australian fishes. VI. Some caligoid species.
Ann. Mag. Nat. Hist., Ser 13, 9: 563-70.
MarsHattr, S. M., and Orr, A. P., 1954——Hatching in Calanus finmarchicus and some
other copepods. J. mar. biol. Ass. U.K., 33: 393-401.
Witson, C. B., 1905.—North American parasitic copepods belonging to the family
Caligidae. Part I. The Caliginae. Proc. U.S. nat. Mus., 28: 479-672.
, 1906.—Report on some parasitic Copepoda collected by Prof. Herdman at
Ceylon, 1902. Rept. Govt. Ceylon Pearl Oyster Fish. Gulf Mannar (Herdman)
Pt. 5: 189-210.
, 1907a.—North American parasitic copepods belonging to the family Caligidae.
Part 2. The Trebinae and Huryphorinae. Proc. U.S. nat. Mus., 31: 669-720.
, 1970b—A revision of the Pandarinae and Cecropinae. Proc. U.S. nat. Mus..
33 : 323-490.
, 191la.—North American parasitic copepods belonging to the family Lernaeo-
podidae. Proc. U.S. nat. Mus., 39: 186-226.
, 19116.—North American parasitic copepods belonging to the family Ergasilidae.
Proc. U.S. nat. Mus., 39: 263-400.
YamacutTi, S., 1963.—‘Parasitic Copepoda and Branchiura of fishes.” New York.
Interscience.
482 ABSTRACT OF PROCEEDINGS
ABSTRACT OF PROCEEDINGS
ORDINARY GENERAL MEETING
27th Marcu, 1968
Professor T. G. Vallance, President, in the chair.
The minutes of the last Ordinary General Meeting (29th November, 1967)
and of the Special General Meeting (29th November, 1967) were taken as read
and signed.
The Chairman announced that the Council had elected the following
Ordinary Members of the Society: Messrs. D. F. Blaxell, D.D.A., B.Sc.,
University of New South Wales, Kensington; and J. R. J. French, B.Sc. (For.),
A.I.W.Sc., Cremorne, N.S.W.
The Chairman announced that library accessions amounting to 53
volumes, 290 parts or numbers, 9 bulletins, 4 reports and 3 pamphlets, total
359, had been received since the last meeting.
The Chairman drew the attention of members to the Australasian Native
Orchid Society, its objects and its publication “Orchadian”; also to the
Australian Research Grants Committee, Department of Education and
Science, Canberra, and its research projects.
PAPERS READ
(By title only, an opportunity for discussion to be given at the April
Ordinary General Meeting)
1. The Rhaphidophoridae (Orthoptera) of Australia. Part 7. A new genus
from the Nullarbor Plain, South-western Australia. By Aola M. Richards.
A new genus is erected, and the new species is described from limestone
caves on the Nullarbor Plain in south-western Australia.
2. A review of the genus Halocynthia Verrill 1879. By Patricia Kott.
The genus Halocynthia Verrill is reduced by synonymy to six closely
related species distinguished only by the condition of the gonads and: the
branchial and atrial spines. Considerable variation in external appearance
is demonstrated within a single species. Species occur in the sublittoral fringe
of land masses and generally have a wide latitudinal range. Their distribution
appears to be limited mainly by deep waters. The genus appears to be an
ancient one and it may represent a relict of the Tethys Sea fauna.
3. The mucosa of the stomach of the wombat (Vombatus hirsutus) with
special reference to the cardiogastric gland. By D. J. Hingson and G. W.
Milton. (Communicated by Dr. Mervyn Griffiths.)
The specialized cardiogastric gland region of wombat stomach which is
characteristic of koala and beaver stomachs as well, is located on the lesser
curve near the oesophageal opening. The cardiogastric gland in the wombat
is distinctive because of its complex group of mucosal sacculations which
open into the stomach lumen via 25 or 30 large crater-like ostia. The mucosa
of this gland contains long, straight, closely packed, unbranched gastric
glands composed of the cell types found elsewhere in the stomach, with chief
cells concentrated at the base of the glands. Parietal cells are present in
great abundance. Typical surface and neck mucous and argyrophilic cells
ABSTRACT OF PROCKEDINGS 483
are also present. The bizarre cardiogastric specialization in the wombat is
thus not cytologically a separate organ from the stomach. However, it does
contribute greatly to the total secretory cell mass of the stomach.
4. The secretory capacity of the stomach of the wombat (Vombatus
hirsutus) and the cardiogastric gland. By G. W. Milton, D. J. Hingson and
K. P. George. (Communicated by Dr. Mervyn Griffiths.)
The secretory capacity of the stomach of the wombat (Vombatus
lirsutus), and of the cardiogastric gland of this animal has been studied.
It was found that the secretory power of this stomach resembles that of man
and animals commonly used in gastric research. The concentrations of ions
in gastric juice generally fitted the Hollander two-component theory of gastric
secretion. A close correlation between the concentration of pepsin and of K*
was demonstrated. The electrophoretic pattern of the gastric juice of the
wombat resembled ‘that obtained from the gastric juice of man. The maximum
secretory capacity of the stomach of this animal was lower than that of man.
It was found that a considerable increase in the gastric output could be
obtained by augmenting the effects of histamine stimulation by injections of
insulin.
5. A taxonomic review of the genus Mixophyes (Anura, Leptodactylidae).
By I. R. Straughan.
Two new species of Mixophyes Gunther are described and the two sub-
species already defined are elevated to species.
OTHER BUSINESS
Dr. I. V. Newman drew attention to signs he had observed of preparations
to lay a sewer line on the surface beside the bed of the Lane Cove River near
its headwaters between Wahroonga and Normanhurst. The steepness of the
valley sides in the uppermost reaches mean great scarring and destruction.
In certain parts of this region is some of the best forest in the Sydney region.
He pointed out that the price of Conservation is Eternal Vigilance.
ORDINARY GENERAL MEETING
24th AprIL, 1968
Professor T. G. Vallance, President, occupied the chair.
The minutes of the last Ordinary General Meeting (27th March, 1968)
were read and confirmed.
The Chairman announced that the Council had elected the following
office-bearers for the 1968-69 session: Vice-Presidents: Mr. L. A. S. Johnson,
Professor R. C. Carolin, Dr. D. T. Anderson and Miss Elizabeth C. Pope;
Honorary Treasurer: Dr. A. B. Walkom; Honorary Secretary: Mr. R. H.
Anderson.
The Chairman announced that the Council had elected Mrs. Elizabeth J.
Hayden, B.Sc. (Melb.)., Burwood, N.S.W., and Dr. A. E. Wood, B.Sc.Agr.,
Ph.D., Bexley, N.S.W., Ordinary Members of the Society.
The Chairman announced that library accessions amounting to 15
volumes, 181 parts or numbers, 7 bulletins, 7 reports and 4 pamphlets, total
214, had been received since the last meeting.
484 ABSTRACT OF PROCEEDINGS
The Chairman drew the attention of members to the notice notifying that
the Australian Research Grants Committee is at present calling applications
for support for research projects in 1969.
The Chairman also announced that no Ordinary General Meeting will be
held in May.
PAPERS READ
1. A new bdellourid-like triclad turbellarian: ectoconsortic on Murray
River Chelonia. By L. R. Richardson.
2. The stratigraphy of the Sofala—Hill End—Euchareena region. By G. H.
Packham.
3. The constitution, distribution and relationships of the Australian
Decapod Crustacea. By D. J. G. Griffin and J. C. Yaldwyn.
4. The embryology of Hpaltes australis Less. (Compositae). By Gwenda
L. Davis.
5. Plants grazed by Red Kangaroos, Megaleia rufa (Desmarest), in
central Australia. By G. Chippendale.
LECTURETTE
Dr. F. H. Talbot, Director of the Australian Museum, Sydney, delivered a
very interesting lecturette, illustrated with colour transparencies, entitled
“Hishes and Corals—ecological results of a study on One Tree Island Reef,
Great Barrier Reef”.
ORDINARY GENERAL MEETING
26th JuNnH, 1968
Professor T. G. Vallance, President, occupied the chair.
The minutes of the last Ordinary General Meeting (24th April, 1968)
were read and confirmed.
The Chairman announced that library accessions amounting to 28
volumes, 453 parts or numbers, 3 bulletins, 11 reports and 47 pamphlets, total
542, had been received since the last meeting.
The Chairman announced that “Australian Natural History’, Vol. 14,
No. 8 (December, 1963) is missing from the Society’s set and is now out of
print. He suggested that some member with a copy might be willing to make
it available to complete the set.
PAPERS READ
1. Aphrophyllum (Rugosa) from Lower Carboniferous limestones near
Bingara, New South Wales. By R. K. Jull. (Communicated by Mr. R. H.
Anderson.)
2. Type specimens in the Macleay Museum, University of Sydney. I. Fishes.
By P. J. Stanbury. (Communicated by Dr. D. T. Anderson.)
3. Replacement name for the preoccupied genus name Odinia Perrier 1885
(Echinodermata : Asteroidea). By A. J. Dartnall, D. L. Pawson, Elizabeth
C. Pope and B. J. Smith.
4. Permian faunas and sediments from the South Marulan district, New
South Wales. By R. E. Wass and I. G. Gould.
es
ABSTRACT. OF PROCHHDINGS 485
HX HIBIT
Mr. G. P. Whitley exhibited two unpublished photographic portraits,
from the Australian Museum’s archives, of naturalists active in Australia over
a century ago. One was of George French Angas (1822-1886) who had been
Secretary of the Australian Museum and is famous for his artistic illustra-
tions; the other of Gerard Krefft (1880-1881), a former Curator and the
zoologist who first described the Queensland lungfish.
LECTURETTE
Miss Elizabeth C. Pope, Australian Museum, Sydney, gave a lecturette
on the “Star fishes of the Fiji reefs”, illustrated by colour transparencies.
ORDINARY GENERAL MEETING
3lst Juty, 1968
Professor T. G. Vallance, President, occupied the chair.
The minutes of the last Ordinary General Meeting (26th June, 1968)
were taken as read and confirmed.
The Chairman announced that Dr. F. H. Talbot, Director of the
Australian Museum, had been elected a member of Council in place of
Professor J. M. Vincent.
The Chairman announced that library accessions amounting to 11
volumes, 138 parts or numbers, 8 bulletins, 5 reports and 3 pamphlets, total
165, had been received since the last meeting.
PAPERS TAKEN AS READ
(By title only, an opportunity for discussion to be given at the
September Ordinary General Meeting) :
1. On the first occurrence of a Climacograptus bicornis with a modified
basal assemblage in Australia. By H. Moors.
2. Chromosome location and linkage studies involving the Pm3 locus for
powdery mildew resistance in wheat. By R. A. McIntosh and E. P. Baker.
3. The vegetation of the Boorabbin and Lake Johnston areas, Western
Australia. By J. S. Beard. (Communicated by Mr. R. H. Anderson.)
ORDINARY GENERAL MEETING
25th SEPTEMBER, 1968
Professor T. G. Vallance, President, occupied the chair.
The minutes of the last Ordinary General Meeting (31st July, 1968)
were read and confirmed.
The Chairman announced that the Council had elected the following
Ordinary Members of the Society: Mr. James Burns, Avalon Beach, 2107;
Miss Estelle M. Canning, Canberra, 2600; Miss Margaret L. Debenham, Strath-
field, 21385; Dr. R. N. Richards, Armidale, 2350; and Dr. Peter Stanbury.
Macleay Museum, University of Sydney, 2006.
486 ABSTRACT OF PROCEEDINGS
The Chairman extended, on behalf of members, congratulations to Mr.
Abdul Khan on his appointment to a lectureship at the new University at
Lahore, Pakistan.
The Chairman announced that library accessions amounting to 19
volumes, 148 parts or numbers, 6 bulletins, 4 reports and 2 pamphlets, total
179, had been received since the last meeting.
The Chairman also announced that the Council is prepared to receive
applications for Linnean Macleay Fellowships tenable for one year from
Ist January, 1969, from qualified candidates. Each applicant must be a
member of this Society and be a graduate in Science or Agricultural Science
of the University of Sydney. The range of actual (tax-free) salary is,
according to qualifications, up to a maximum of A$3,200 per annum. Applica-
tions should be lodged with the Honorary Secretary, who will give further
details and information, not later than Wednesday, 6th November, 1968.
The Chairman drew the attention of members to a “Symposium on Arid
Lands” sponsored by the Australian Academy of Science, Canberra, on
19-21 May, 1969.
The Chairman drew the attention of members to an invitation by the
Wildlife Preservation Society of Australia to attend a function on 6th
November in honour of Mr. Allen Strom.
PAPERS READ
1. A study of some smuts of Hehinochloa spp. By R. A. Fullarton and
R. F. N. Langdon.
2. A viviparous species of Patiriella (Asteroidea, Asterinidae). from
Tasmania. By A. J. Daritnall.
3. The nasal mites of Queensland birds (Acari : Dermanyssidae,
Ereynetidae and Epidermoptidae). By R. Domrow.
LECTURETTE
An illustrated lecturette entitled “Some new aspects of Crustacean
development” was delivered by Dr. D. T. Anderson, Department of Zoology,
University of Sydney.
ORDINARY GENERAL MEETING
30th OcrossEr, 1968
Professor T. G. Vallance, President, in the chair.
The minutes of the last Ordinary General Meeting (25th September, 1968)
were read and confirmed.
The Chairman, on behalf of members, extended congratulations to Dr.
Beryl Nashar on her appointment as Dean of the Faculty of Science,
Newcastle University.
The Chairman announced that Miss Christine D. Clarke, Milson’s Point,
and Mr. L. A. Nielsen, Jandowae, Queensland, had been elected by the Council
to membership of the Society.
The Chairman announced that library accessions amounting to 13
volumes, 162 parts or numbers, 8 bulletins, 8 reports and 3 pamphlets, total
194, had been received since the last meeting.
ABSTRACT OF PROCHEDINGS 487
The Chairman announced that the Council is prepared to receive appli-
cations for Linnean Macleay Fellowships tenable for one year from Ist
January, 1969, from qualified candidates.
PAPERS RHAD
1. Type specimens in the Macleay Museum, University of Sydney. II.
Amphibians and reptiles. By Judy Goldman, L. Hill and P. J. Stanbury.
2. Notes on Vittadinia triloba sens. lat. (Compositae). By Nancy T.
Burbidge. (Communicated by Dr. Joyce W. Vickery.)
LECTURETTR
An illustrated lecturette entitled “Some local Charophytes” was delivered
by Dr. A. T. Hotchkiss, Department of Botany, University of Sydney.
ORDINARY GENERAL MEETING
27th Novemser, 1968
Professor T. G. Vallance, President, in the chair.
The minutes of the last Ordinary General Meeting (30th October, 1968)
were read and confirmed.
The Chairman announced that the following had been elected by the
Council to membership of the Society: Mr. D. R. Goodfellow, Carlingford,
N.S.W.; Mr. J. H. Phippard, B.Pharm., University of Queensland, Brisbane,
Queensland; Dr. C. J. Quinn, B.Se.(Hons.), Ph.D., University of New South
Wales, Kensington, N.S.W., and Mr. B. V. Timms, B.Sc.(Hons.), Avondale
College, Cooranbong, N.S.W.
The Chairman announced that the Council had re-appointed Miss Alison
K. Dandie, B.Sc.(Hons.), to a Linnean Macleay Fellowship in Botany for
one year from ist January, 1969.
The Chairman referred to the death on 2nd November, 1968, of Sir
Harold Raggatt, a distinguished geologist, who had been a member of the
Society since 1929.
The Chairman also announced that library accessions amounting to
36 volumes, 253 parts or numbers, 10 bulletins, 5 reports and 2 pamphlets,
total 306, had been received since the last meeting.
PAPERS RHEAD
1. A revision of the family Agnesiidae Huntsman 1912, with particular
reference to Agnesia glaciata Michaelsen 1898. By Patricia Kott.
2. Type specimens in the Macleay Museum, University of Sydney. III.
Birds. By P. J. Stanbury.
3. Type specimens in the Macleay Museum, University of Sydney. IV.
Mammals. By P. J. Stanbury.
4. Hatching and larval development of a species belonging to the
Copepod family, Ascidicolidae, a parasite of the ascidian, Styela etheridgii
Herman. By D. T. Anderson and G. T. Rossiter.
5. Hatching and larval development of Dissonus nudiventris Kabata
(Copepoda, Fam. Dissonidae), a gill parasite of the Port Jackson shark.
By D. T. Anderson and G. T. Rossiter.
488 ABSTRACT OF PROCEEDINGS
NOTES AND EXHIBITS
Mr. G. P. Whitley exhibited, and commented upon, some Japanese
papers on poisonous crabs, a topic which had been discussed at last August’s
Seminar on ichthyosanotoxism arranged by the South Pacific Commission
at Tahiti. x
Dr. A. Hotchkiss discussed the ripening and after-ripening in fruits
of Ruppia spiralis Dumort. The fruits of Ruppia spiralis were shown in
photographs illustrating several stages in ripening. At the stage of maturity
on the plant the fruits are covered with smooth, green, soft outer tissues.
The fruits are immediately deciduous and undergo a period of after-ripening
for about two weeks during which the inner pericarp hardens and darkens,
the outer tissues die, decay and eventually are sloughed off both the fruit
and the stipe. The fruit at this stage is terminated by a slender beak, and
its surface is covered with stout spines composed of thick-walled, irregularly
branched sclereids projecting from the outer surface of the pericarp wall.
At germination, a calyptra bends back from the convex side, pushed outward
by the emerging radical.
The President, Professor T. G. Vallance, referred to the centenary (1967)
of the Rev. W. B. Clarke’s work “Remarks on the Sedimentary Formations
of New South Wales”. From a summary statement some 15 pages in length
the work grew in the space of eleven years (and four editions) to become a
book of 165 pages, completed only a fortnight before Clarke’s death in
June, 1878. Clarke has been called the “Father of Australian Geology”, and
this series of editions provides a means of examining the development of
his ideas and attitudes on Australian stratigraphy. Close, comparative study
of these documents would amply repay the attention of some historian
of geology.
Some examples of Clarke’s developing stratigraphical synthesis, such
as his increasing confidence in the existence of Devonian strata and doubts
as to the ages of the upper part of the coal sequence and the strata above
it in the Sydney basin, were briefly discussed.
Copies of the four editions were exhibited. Brief details of these were
listed because the first three issues, at least, are not readily traceable.
Ist edition—pp. 65-80 in Catalogue of the Natural and Industrial
Products of New South Wales forwarded to the Paris Universal Exhibition
of 1867, by the New South Wales Exhibition Commissioners. Government
Printer, Sydney, 1867.
2nd edition—pp. 505-531 in Industrial Progress of New South Wales:
being a report of the Intercolonial Exhibition of 1870, at Sydney; together
with a variety of papers illustrative of the industrial resources of the colony.
Government Printer, Sydney, 1871.
3rd edition— pp. 149-206 in New South Wales Intercolonial and
Philadelphia International Exhibition. Mines and Mineral Statistics of New
South Wales, and notes on the geological collection of the Department of
MAIVES Sia. ice siete Government Printer, Sydney, 1875.
4th edition—Remarks on the Sedimentary Formations of New South
Walesabs SHO. Bee Government Printer, Sydney, 1878.
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N
LIST OF MEMBERS
(15th December, 1968)
ORDINARY MEMBERS
(An asterisk ( * ) denotes Life Member)
Abbie, Professor Andrew Arthur, M.D., B.S., B.Se., Ph.D., c/- University of Adelaide,
Adelaide, South Australia, 5000.
*Allman, Stuart Leo, B.Sc.Agr., M.Se., 99 Cumberland Avenue, Collaroy, N.S.W., 2097.
Anderson, Derek John, Ph.D., School of Biological Sciences, Botany Building, Sydney
University, 2006.
Anderson, Donald Thomas, B.Sc., Ph.D., School of Biological Sciences, Department of
Zoology, Sydney University, 2006.
Anderson, Mrs. Jennifer Mercianna Elizabeth, B.Sc.Agr., 51 Ocean Street, Woollahra,
N.S.W., 2025.
Anderson, Robert Henry, B.Sc.Agr., 19 Kareela Road, Chatswood, N.S.W., 2067.
Andrew, Mrs. Phillipa Audrey, M.Sc. (née Croucher), 10 Black Street, Watsonia, Victoria,
3087. .
Ardley, John Henry, B.Sc. (N.Z.), Messrs. William Cooper and Nephews (Australia) Pty.
Ltd., P.O. Box 12, Concord, N.S.W., 2137.
*Armstrong, Jack Walter Trench, “* Cullingera’”’, Nyngan, N.S.W., 2825.
Ashton, David Hungerford, B.Sc., Ph.D., 92 Warrigal Road, Surrey Hills, Victoria, 3127.
Aurousseau, Marcel, B.Sc., 229 Woodland Street, Balgowlah, N.S.W., 2093.
Bain, Miss Joan Maud, M.Se., Ph.D., 18 Onyx Road, Artarmon, N.S.W., 2064.
Baker, Professor Eldred Perey, B.Se.Agr., Ph.D., Department of Agricultural Botany,
Sydney University, 2006.
Ballantyne, Miss Barbara Jean, B.Sc.Agr., N.S.W. Department of Agriculture, Private
Mail Bag No. 10, Rydalmere, N.S.W., 2116.
Bamber, Richard Kenneth, F.S.T.C., 113 Lucinda Avenue South, Wahroonga, N.S.W.,
2076.
*Barber, Professor Horace Newton, M.S., Ph.D., F.A.A., School of Biological Sciences,
Department of Botany, University of N.S.W., P.O. Box 1, Kensington, N.S.W., 2033.
Barlow, Bryan Alwyn, B.Sc., Ph.D., School of Biological Sciences, The Flinders University
of South Australia, Bedford Park, South Australia, 5042.
Basden, Ralph, M.Ed., B.Sc. (Lond.), F.R.A.C.1., A.S.T.C., 183 Parkway Avenue,
Hamilton, N.S.W., 2303.
Batley, Alan Francis, A.C.A., 123 Burns Road, Wahroonga, N.S.W., 2076.
Baur, George Norton, B.Sc., B.Se.For., Dip.For., 3 Mary Street, Beecroft, N.S.W., 2119.
*Beadle, Professor Noel Charles William, D.Sc., University of New England, Armidale,
N.S.W., 2350.
Bearup, Arthur Joseph, B.Sc., 66 Pacific Avenue, Penshurst, N.S.W., 2222.
Beattie, Joan Marion, D.Sc. (née Crockford), 2 Grace Avenue, Beecroft, N.S.W., 2119.
Bedford, Geoffrey Owen, B.Sc., 87 Jacob Street, Bankstown, N.S.W., 2200.
Bennett, Miss Isobel Ida, Hon.M.Se., School of Biological Sciences, Department of
Zoology, Sydney University, 2006.
Bertus, Anthony Lawrence, B.Sc., Biology Branch, N.S.W. Department of Agriculture,
Private Mail Bag, No. 10, Rydalmere, N.S.W., 2116.
Besly, Miss Mary Ann Catherine, B.A., School of Biological Sciences, Department of
Zoology, Sydney University, 2006.
Bishop, James Arthur, Department of Genetics, The University of Liverpool, Liverpool 3,
England.
Blackmore, John Allan Philip, LL.B. (Syd. Univ.), 25 Holden Street, Ashfield, N.S.W.,
2131. ;
Blake, Clifford Douglas, B.Se.Agr., Ph.D., Faculty of Agriculture, Sydney University,
2006.
Blake, Stanley Thatcher, D.Sc. (Q’ld.), 1110 Waterworks Road, The Gap, Queensland, 4061.
Blaxell, Donald Frederick, D.D.A., B.Sc., School of Biological Sciences, University of New
South Wales, P.O. Box 1, Kensington, N.S.W., 2033.
Bourke, Terrence Victor, B.Sc.Agr., c/- Department of Agriculture, Stock and Fisheries,
Popondetta, Papua.
Boyd, Robert Alexander, B.Se., Department of Botany, University of New England,
Armidale, N.S.W., 2350.
Brett. Robert Gordon Lindsay, B.Sc., 48 Main Road, Lindisfarne, Tasmania, 7015.
Brewer, Ilma Mary, D.Sc., 13 Wentworth Road, Vaucluse, N.S.W., 2030.
Briggs, Miss Barbara Gillian, Ph.D., National Herbarium of N.S.W., Royal Botanic
Gardens, Sydney, 2000.
Browne, Ida Alison, D.Sc. (née Brown), 363 Edgecliff Road, Edgecliff, N.S.W., 2027.
Browne, William Rowan. D.Sc., F.A.A., 363 Edgecliff Road, Edgecliff, N.S.W., 2927.
Burden, John Henry, 1 Havilah Street, Chatswood, N.S.W., 2067.
490.
1931
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LIST OF MEMBERS
*Burges, Professor Norman Alan, M.Sc., Ph.D., Professor of Botany, University of Liver-
pool, Liverpool, England.
Burgess, The Rev. Colin E. B. H., Parks and Gardens Section, Department of the
Interior, Canberra, A.C.T., 2600.
Burgess, Ian Peter, B.Sc.For., Dip.For., The Forestry Office, Coff’s Harbour, N.S.W.,
2450. ‘
Burns, James, A.A.S.A., 127 Plateau Road, Avalon Beach, N.S.W., 2107.
Cady, Leo Isaac, P.O. Box 88, Kiama, N.S.W., 2533.
Campbell, Keith George, D.F.C., B.Sc.For., Dip.For., M.Sc., 17 Third Avenue, Epping,
N.S.W., 2121.
Campbell, Thomas Graham, Division of Entomology, C.S.I.R.O., P.O. Box 109, City,
Canberra, A.C.T., 2601.
Canning, Miss Estelle Margaret, B.Sc. (Melb.), c/- Canberra Botanic Gardens, Parks and
Gardens Branch, Department of Interior, Canberra, A.C.T., 2600.
*Carey, Professor Samuel Warren, D.Sc., Geology Department, University of Tasmania,
Hobart, Tasmania, 7000.
Carne, Phillip Broughton, B.Agr.Sci. (Melb.), Ph.D. (London), D.I.C., C.S.I.R.O., Division
of Entomology, P.O. Box 109, City, Canberra, A.C.T., 2601.
Carolin, Professor Roger Charles, B.Sc., A.R.C.S., Ph.D., School of Biological Sciences,
Department of Botany, Sydney University, 2006.
Casimir, Max, B.Sc.Agr., Entomological Branch, N.S.W. Department of Agriculture,
Private Mail Bag, No. 10, Rydalmere, N.S.W., 2116.
*Chadwick, Clarence Earl, B.Sc., Entomological Branch, N.S.W. Department of Agri-
culture, Private Mail Bag No. 10, Rydalmere, N.S.W., 2116.
Chambers, Thomas Carrick, M.Sc. (N.Z.), Ph.D., Botany School, University of Melbourne,
Parkville, Victoria, 3052.
Child, John, M.A., B.Comm. (N.Z.), D.Phil. (Oxon.), Department of Economics, Otago
University, Box 56, Dunedin, New Zealand.
Chippendale, George McCartney, B.Sc., 4 Raoul Place, Lyons, A.C.T., 2606.
Christian, Stanley Hinton, Malaria Research Unit and School, Kundiawa, Eastern High-
lands, Territory of Papua and New Guinea.
*Churchward, John Gordon, B.Sc.Agr., Ph.D., “ Erlangga’”’, Glen Shian Lane, Mount
Eliza, Victoria, 3930.
Clark, Laurance Ross, M.Sc., e/- C.S.I.R.O., Division of Entomology, P.O. Box 109, City,
Canberra, A.C.T., 2601.
Clarke, Miss Christine Dorothea, B. Sce.(Hons.), 26a Alfred Street, Milson’s Point, N.S.W.,
2061.
Clarke, Miss Lesley Dorothy, Ph.D., 4 Gordon Crescent, Eastwood, N.S.W., 2122.
Clarke, Mrs. Muriel Catherine, M.Sc. (née Morris), 122 Swan Street, Morpeth, N.S.W.
232%
Cleland, Professor Sir John Burton, M.D., Ch.M., C.B.E., 1 Dashwood Road, Beaumont,
Adelaide, South Australia, 5066.
Clough, Barry Francis, B.Sc.Agr., School of Biological Sciences, Department of Botany,
Sydney University, 2006.
Clyne, Mrs. Densey, 7 Catalpa Crescent, Turramurra, N.S.W., 2074.
Cogger, Harold George, M.Sc., Department of Zoology, Macquarie University, North
Ryde, N.S.W., 2113. ;
Colless, Donald Henry, Ph.D. (Univ. of Malaya). c/- Division of Entomology, C.S.I.R.O.,
P.O. Box 109, City, Canberra, A.C.T., 2601.
Common, Jan Francis Bell, M.A., M.Sc.Agr., C.S.I.R.O., Division of Entomology, P.O.
Box 109, City, Canberra, A.C.T., 2601.
Conroy, Brian Alfred, International House, Sydney University, 2006.
Copland, Stephen John, M.Sc., 15 Chilton Parade, Warrawee, N.S.W., 2074.
Costin, Alex Baillie, B.Se.Agr., C.S.I.R.O., Division of Plant Industry, P.O. Box 109,
City, Canberra, A.C.T., 2601.
Craddock, Miss Elysse Margaret, 36 Lyons Road, Drummoyne, N.S.W., 2047.
Crawford, Lindsay Dinham, B.Sc., ec/- Victorian Plant Research Institute, Department
of Agriculture, Burnley Gardens, Melbourne, Victoria, 3000.
Crook, Keith Alan Waterhouse, M.Sc., Ph.D. (New England), Department of Geology,
Australian National University, G.P.O. Box 197, Canberra, A.C.T., 2601.
Dandie, Miss Alison Kay, B.Sec.(Hons.), Dip.Ed., 69 Waitara Parade, Hurstville, N.S.W.,
N.S.W.. 2220.
Dart, Peter John, B.Sc.Agr., Ph.D., Soil Microbiology Department, Rothamsted Experi-
mental Station, Harpenden, Herts., England.
Dartnall, Alan John, B.Sc., 7 Forbes Avenue, West Hobart, Tasmania, 7000.
Davies, Stephen John James Frank, B.A. (Cantab.), Ph.D., C.S.I.R.O., Private Bag,
Nedlands, Western Australia, 6009.
Davis, Professor Gwenda Louise, Ph.D., B.Sc., Faculty of Science, University of New
England, Armidale, N.S.W., 2350.
Debenham, Miss Margaret Lee, B.Sc., 42 Hunter Street, Strathfield, N.S.W., 2135.
1953
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LIST OF MEMBERS 49]
De Nardi, Mrs. Jan Christina, B.Sc. (Q’ld.) (née Morrow), 6/81 New South Head Road,
Vaucluse, N.S.W., 2030.
Dobrotworsky, Nikolai V., M.Sc., Ph.D., Department of Zoology, University of Melbourne,
Parkville, Victoria, 3052.
Domrow, Robert, B.A., B.Sc., Queensland Institute of Medical Research, Herston Road,
Herston, Queensland, 4006.
Dorman, Herbert Clifford, J.P., A.S.T.C. (Dip.Chem.), Dip.Soc.Stud. (Sydney), Rodgers
Street, Teralba, N.S.W., 2284.
Douglas, Geoffrey William, B.Agr.Se., Deputy Chairman, Vermin and Noxious Weeds
Destruction Board, Department of Crown Lands and Survey, Treasury Place,
Melbourne, Victoria, 3002.
Durie, Peter Harold, M.Sc., C.S.I.R.O., Veterinary Parasitology Laboratory, Yeerongpilly,
Queensland, 4105.
Dyce, Alan Lindsay, B.S¢.Agr., 48 Queen’s Road, Asquith, N.S.W., 2078.
Edwards, Dare William, B.Se.Agr., Forestry Commission of N.S.W., Division of Wood
Technology, 96 Harrington Street, Sydney, 2000.
Edwards, Edward John, B.A., B.Se., Dip.Ed., 38 Shirlow Avenue, Faulconbridge, N.S.W..,
2776. ‘
Endean, Robert, M.Se., Ph.D., Department of Zoology, University of Queensland, St.
Lucia, Queensland, 4067. -
English, Miss Kathleen Mary Isabel, B.Sc., 6/168 Norton Street, Leichhardt, N.S.W., 2040.
Evans, Miss Gretchen Pamela, M.Sc., 27 Frederick Street, Taringa, Queensland, 4066.
Facer, Richard Andrew, Department of Geology, Wollongong University College
Wollongong, N.S.W., 2500.
*Fairey, Kenneth David, Box 1176, G.P.O., Sydney, 2001.
Filewood, Lionel Winston Charles, ¢c/- Department of Agriculture, Stock and Fisheries,,
KKonedobu, Papua.
Florence, Ross Garth. M.Sc.For., Ph.D., The Australian National University, Department
of Forestry, P.O. Box 4, Canberra, A.C.T., 2600.
Ford, Miss Judith Helen, 18 Central Avenue, Mosman, N.S.W., 2088.
Fraser, Miss Lilian Ross, D.Sec., 1 Laurence Street, Pennant Hills, N.S.W., 2120.
French, John Richard Joseph, B.Sc.(For.), A.I.W.Sc., 5 Brierley Street, Cremorne, N.S.W.,
2090.
*Garretty, Michael Duhan, D.Sc.. Box 763, Melbourne, Victoria, 3001.
Greenwood, William Frederick Neville, 11 Wentworth Avenue, Waitara, N.S.W., 2977.
Griffin, David Michael, M.A., Ph.D. (Cantab.), School of Agriculture, Sydney University,
2006.
*Griffiths, Mrs. Mabel, B.Sc. (née Crust), 54 Delmar Parade, Dee Why, N.S.W., 2099.
Griffiths, Mervyn Edward, D.Se., Wildlife Survey Section, C.S.I.R.O., P.O. Box 109,
City, Canberra, A.C.T., 2601.
*Gunther, Carl Ernest Mitchelmore, M.B., B.S., D.T.M., D.T.M. & H. (England), M.B.E.,
29 Flaumont Avenue, Lane Cove, N.S.W., 2066.
Hadlington, Phillip Walter. B.Sc.Agr., 129 Condamine Street, Balgowlah, N.S.W., 2093.
Hannon, Miss Nola Jean, B.Se., Ph.D., 22 Leeder Avenue, Penshurst, N.S.W.. 2222.
Harden, Mrs. Gwenneth Jean, M.Sc. (née Hindmarsh), Kellys Plains Road, Armidale,
N.S.W., 2350.
Hardwick, Reginald Leslie, B.Sc., 183 Richmond Road, Kingswood, N.S.W., 2750.
Hartigan, Desmond John, B.Sc.Agr., 75 Northwood Road, Northwood, N.S.W., 2066.
Hayden, Mrs. Elizabeth Jean, B.Sc. (Melb.), Botany Department, Australian National
University, P.O. Box 4, Canberra, A.C.T., 2600.
Hennelly, John Patten Forde, B.Se., Highs Road, West Pennant Hills, N.S.W., 2120.
Hewitt, Bernard Robert, B.A. (Q’ld.), B.Se. (Syd.), M.Se. (N.S.W.), A.R.A.C.I., Senior
Lecturer in Chemistry, University of Malawi, Limbe, Malawi, Africa.
Hewson, Miss Helen Joan, B.Sc.(Hons.), Ph.D., Department of Botany, School of General
Studies, Australian National University, P.O. Box 4, Canberra, A.C.T., 2600.
Higginson, Francis Ross, B.Sc.Agr.(Hons.), Ph.D., Soil Conservation Service of N.S.W.,
Box 4293, G.P.O., Sydney, 2001.
Hill, Miss Dorothy, M.Sc., Ph.D., Department of Geology, University of Queensland,
Brisbane, Queensland, 4067. :
*Hindmarsh, Miss Mary Maclean, B.Sc., Ph.D., 4 Recreation Avenue, Roseville, N.S.W.,
2069.
*Holder, Miss Lynette Anne, B.Sc., 48 Rutledge Street, Eastwood, N.S.W., 2122.
Holland, Ray James Thurstan, M.A. (Syd.), M.A.C.E., e/- Sydney Grammar School,
College Street, Sydney, 2000.
1953 *Hotchkiss, Professor Arland Tillotson, M.S., Ph.D. (Cornell), Department of Biology, Uni-
1956
versity of Louisville, Louisville, Kentucky, 40208, U.S.A.
*Hotchkiss, Mrs. Doreen Elizabeth, Ph.D., B.A., M.A. (née Maxwell), 2440 Longest Avenue,
Louisville, Kentucky, 40208, U.S.A.
492 -
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LIST OF MEMBERS
Hoult, Errol Hubert, B.Sc.(Hons.), Department of Botany, University of New England,
Armidale, N.S.W., 2350.
Humphrey, George Frederick, M.Sc., Ph.D., C.S.I.R.O. Marine Biological Laboratory,
Box 21, Cronulla, N.S.W., 2230.
Ingram, Cyril Keith, B.A., B.Ec., Office df Inspector of Schools, 2/306 Malton Road,
Epping North, N.S.W., 2121.
Jackes, Mrs. Betsy Rivers, B.Sc., Ph.D. (Univ. Chicago) (née Paterson), 21 Nebo Road,
Mackay, Queensland, 4740. ;
Jacobs, Miss Janice Lorraine, B.Sc., School of Biological Sciences, Department of Botany,
' Sydney University, 2006.
Jacobs, Maxwell Ralph, D.Ing., M.Se., Dip.For., 32 Arthur Circle, Forrest, A.C.T., 2603.
Jacobs, Surrey Wilfred Laurence, 7 Yarrara Road, Pymble, N.S.W., 2073.
Jacobson, Miss Constance Mary, M.Sc., Ph.D., School of Biological Sciences, Department
of Zoology, Sydney University, 2006.
James, Sidney Herbert, M.Sc., 54 Holmfirth Street, Mt. Lawley, Western Australia, 6050.
Jancey, Robert Christopher, M.Sc., Ph.D., e/- Department of Botany, University of Western
Ontario, London, Ontario, Canada.
Jefferies, Mrs. Lesly Joan, 14 Denman Street, Hurstville, N.S.W., 2220.
Jenkins, Thomas Benjamin Huw, Ph.D., Department of Geology and Geophysics, Sydney
University, 2006.
Jessup, Rupert William, M.Se., 6 Penno Parade North, Belair, South Australia 5052.
Johnson Lawrence Alexander Sidney, B.Sc.. c/- National Herbarium, Royal Botanic
Gardens, Sydney, 2000.
Johnston, Arthur Nelson, B.Sc.Agr., 224 Greville Street, Chatswood, N.S.W., 2067.
Jolly, Violet Hilary, M.Se., Ph.D., Metropolitan Water, Sewerage and Drainage Board,
P.O. Box 2, Guildford, N.S.W., 2161.
Jones, Edwin Llewelyn, B.A., P.O. Box 196, Leeton, N.S.W., 2705.
Jones, Leslie Patrick, Department of Animal Husbandry, Sydney University, 2000.
Joplin, Miss Germaine Anne, B.A., Ph.D., D.Sc., Department of Geophysics, Australian
National University, Canberra, A.C.T., 2600.
Judd, Howard Kenniwell, Minnamurra Falls Forest Reserve, Box 14, P.O., Jamberoo,
N.S.W., 2533.
Keast, James Allen, M.Sc., M.A., Ph.D. (Harvard), Professor of Vertebrate Zoology,
Queen’s University, Kingston, Ontario, Canada.
Kerr, Harland Benson, B.Sc.Agr., Ph.D., Summer Institute of Linguistics, P.O. Ukarumpa,
E.H.D., Territory of New Guinea.
Kesteven, Geoffrey Leighton, D.Se., ¢/- Division of Fisheries and Oceanography
C.S.1I.R.0., P.O. Box 21, Cronulla, N.S.W., 2230.
Khan, Abdul Ghaffar, M.Sc., Botany Department, Panjab -University, New-Campus,
Lahore, West-Pakistan.
Kindred, Miss Berenice May, B.Sc., Fachbereich Biologie, Universitat Konstanz, 775
Konstanz, Postfach 733, Germany.
Langdon, Raymond Forbes Newton, M.Agr.Sc., Ph.D., Department of Botany, University
of Queensland, St. Lucia, Queensland, 4067.
Lanyon, Miss Joyce Winifred, B.Sc., Dip.Ed., 35 Gordon Street, Hastwood, N.S.W., 2122.
Lawson, Albert Augustus, ““ Rego House 5 23-25 Foster Street, Sydney, 2000.
Lee, Mrs. Alma Theodora, M.Sc. (née Melvaine), Manor Road, Hornsby. N.S.W.. 2077.
Lee, Professor David Joseph, B.Se., School of Public Health and Tropical Medicine, Sydney
University, 2006.
Littlejohn, Murray John, B.Se., Ph.D. (W.A.), Department of Zoology, University of
Melbecurne, Parkville, Victoria, 3052.
Lothian, Thomas Robert Noel, Botanic Gardens, Wacinide: South Australia, 5000.
Lovedee, Miss Lois Jacqueline, B.Sc. (A.N.U.), 11 Dundilla Road, French’s Forest, N.S.W.,
2086.
Luig, Norbert Harold, Ph.D., c/- Faculty of Agriculture, Sydney University, 2006.
Lyne, Arthur Gordon, B.Se., Ph.D., C.S.I.R.O., Ian Clunies Ross Animal Research
Laboratory, P.O. Box 144, Parramatta, N.S.W., 2150.
Macdonald, Colin Lewis, 7 Watford Close, North Epping. N.S.W., 2121.
Macintosh, Professor Neil William George, M.B., B.S., Department of Anatomy, Sydney
University, 2006.
Mackay, Miss Margaret Muriel, B.Sc.(Hons.), M.Sce., M.I.Biol., J.P., 6 Woodford Street,
Longueville, N.S.W., 2066.
Mackerras, Ian Murray, M.B., Ch.M., B.Sc., C.S.1.R.O., Division of Entomology, P.O. Box
109, City, Canberra, A.C.T., 2601.
*Mair, Herbert Knowles Charles, B.Sc., Royal Botanic Gardens, Sydney, 2000.
Marks, Miss Elizabeth Nesta, M.Sc., Ph.D., Department of Entomology, University of
Queensland, Brisbane, Queensland, 4067.
LIST OF MEMBERS 493
Martin, Angus Anderson, B.Sc. (Hons.), Rand, Department of Zoology, University of
Melbourne, Parkville, Victoria, 3052.
Martin, Anthony Richard Henry, M.A., Ph.D., School of Biological Sciences, Department
of Botany, Sydney University, 2006.
Martin, Mrs. Hilda Ruth Brownell, B.Sc. (née Simons), c/- Mrs. H. I. Simons, 43 Spencer
Road, Killara, N.S.W., 2071.
Martin, Peter Marcus, M.Sc.Agr., Dip.Ed., School of Biological Sciences, Department of
Botany, Sydney University, 2006.
Mather, Mrs. Patricia (née Kott), M.Se., Ph.D. (W.A. and Q’ld), Department of Zoology,
University of Queensland, St. Lucia, Queensland, 4067.
McAlpine, David Kendray, M.Se., 12 St. Thomas Street, Bronte, N.S.W., 2024.
McCulloch, Robert Nicholson, M.B.E., D.Sc.Agr., B.Sec., Cattle Tick Research Station,
Wollongbar, N.S.W., 2480.
*McCusker, Miss Alison, M.Se., Botany Department, University College, Box 9184, Dar es
Salaam, Tanzania.
McDonald, Miss Patricia M., B.Se., Dip.Ed., 33 Holdsworth Street, Neutral Bay, N.S.W.,
2089.
McGarity, John William, M.Se.Agr., Ph.D., Agronomy Department, School of Rural
Science, University of New England, Armidale, N.S.W., 2350.
McGillivray, Donald John, B.Se.For. (Syd.), Dip.For. (Canb.), c/- Royal Botanic Gardens,
Kew, Richmond, Surrey, England.
Mckee, Hugh Shaw, B.A., D.Phil. (Oxon.), Service des Eaux et Foréts, B.P. 285, Noumea,
New Caledonia.
McKenna, Nigel Reece, Department of Education, Konedobu, Papua.
Mercer, Professor Frank Verdun, B.Sc., Ph.D. (Camb.), School of Biological Sciences,
Macquarie University, North Ryde, N.S.W., 2113.
Messmer, Mrs. Pearl Ray, 64 Treatts Road, Lindfield, N.S.W., 2070.
*Meyer, George Rex, B.Sc., Dip.Ed., B.A., M.Ed., Centre for Advancement of Teaching,
Macquarie University, North Ryde, N.S.W., 2113.
*Miller, Allen Horace, B.Sc., Dip.Ed., 89 Kentucky Street, Armidale, N.S.W., 2350.
Millerd, Miss Alison Adéle, Ph.D., C.S.I.R.O., Division of Plant Industry, P.O. Box 109,
City, Canberra, A.C.T., 2601.
Millett, Mervyn Richard Oke, B.A., “ Beeyung”’, 72 McNicol Road, Tecoma, Victoria,
3160
Milward, Norman Edward, B.Sc. (Hons.), M.Sec., Department of Zoology, University
College of Townsville, Pimlico, Townsville, Queensland, 4810.
Moore, Barry Philip, B.Se., Ph.D., D.Phil., C.S.I.R.O., Division of Entomology, P.O. Box
109, City, Canberra, A.C.T., 2601.
Moore, Kenneth Milton, Cutrock Road, Lisarow, N.S.W., 2251.
Moore, Raymond Milton, D.Sc.Agr., 94 Arthur Circle, Forrest, A.C.T., 2603.
Moors, Henry Theodore, B.Se., Department of Geology and Mineralogy, University of
Melbourne, Parkville, Victoria, 3052.
Morgan, Mrs. Eva, M.Sc., 5317 Borland Road, Los Angeles, California, 90032, U.S.A.
Morrison, Gordon Cyril, 34 Leuna Avenue, Wahroonga, N.S.W., 2076.
Moss, Francis John, 37 Avenue Road, Mosman, N.S.W., 2088.
Muirhead, Warren Alexander, B.Sc.Agr., C.S.I.R.O., Irrigation Research Station, Private
Mail Bag, Griffith, N.S.W., 2680.
Mungomery, Reginald William, c/- Bureau of Sugar Experiment Stations, 99 Gregory
Terrace, Brisbane, Queensland, 4000.
Murray, David Ronald, B.Sc., 14 Consul Road, Brookvale, N.S.W., 2100.
Nashar, Professor Beryl, B.Sc., Ph.D., Dip.Ed. (née Scott), 43 Princeton Avenue,
Adamstown Heights, N.S.W., 2289.
Newman, Ivor Vickery, M.Se., Ph.D., F.R.M.S., F.L.S., School of Biological Sciences,
Macquarie University, North Ryde, N.S.W., 2113.
Nicholls, Anthony Oldham, B.Se., School of Botany, University of Melbourne, Parkville,
Victoria, 3052.
Nicholson, Alexander John, C.B.E., D.Sc., F.R.E.S., C.S.I.R.O., Box 109, City, Canberra,
A.C.T., 2601.
Nielsen, Lloyd Alwyn, P.O. Box 12, Jandowae, Queensland, 4410.
*Noble, Norman Scott, D.Sc.Agr., M.Se., D.I.C., Unit 21, 1 Lauderdale Avenue, Fairlight,
N.S.W., 2094.
Noble, Robert Jackson, B.Sc.Agr., Ph.D., 324 Middle Harbour Road, Lindfield, N.S.W.,
2070.
O’Farrell, Professor Antony Frederick Louis, A.R.C.Se., B.Se., F.R.E.S., Department of
Zoology, University of New England, Armidale, N.S.W., 2350.
O’Gower, Professor Alan Kenneth, M.Sec., Ph.D., 20 Gaerloch Avenue, Bondi, N.S.W., 2026.
Osborn, Professor Theodore George Bentley, D.Se., F.L.S.. 34 Invergowrie Avenue,
Highgate, South Australia, 5063.
Oxenford, Reginald Augustus, B.Sc., 107 Alice Street, Grafton, N.S.W., 2460.
494
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LIST OF MEMBERS
Packham, Gordon Howard, B.Sc., Ph.D., Department of Geology, Sydney University,
2006.
*Pasfield, Gordon, B.Sc.Agr., 20 Cooper Street, Strathfield, N.S.W., 2135.
Patrick, John William, B.Sc.Ag.(Hons.), School of Biological Sciences, Macquarie
University, North Ryde, N.S.W., 2113.
Payne, William Herbert, A.S.T.C., A.M.I.E.Aust., M.A.P.E., 250 Picnic Point Road,
Pienie Point, N.S.W., 2213. r
Peacock, William James, B.Sc., Ph.D., C.S.1.R.O., Division of Plant Industry, P.O. Box
109, Canberra City, A.C.T., 2601.
Pedder, Alan Edwin Hardy, M.A. (Cantab.), Institute of Sedimentary and Petroleum
Geology, 3303 33rd Street, N.W. Calgary 44, Alberta, Canada.
Perkins, Frederick Athol, B.Sc.Agr., 93 Bellevue Terrace, Clayfield, Queensland, 4011.
Philip, Graeme Maxwell, M.Sc. (Melb.), Ph.D. (Cantab.), F.G.S., Department of Geo ogy,
University of New England, Armidale, N.S.W., 2350.
Phillips, Miss Marie Elizabeth, M.Sc., Ph.D., Parks and Gardens Section, Department of
the Interior, Canberra, A.C.T., 2600.
Pope, Miss Elizabeth Carington, M.Sc., C.M.Z.S., Australian Museum, P.O. Box A285,
Sydney South, N.S.W., 2000.
Pryor, Professor Lindsay Dixon, M.Sc., Dip.For.. Department of Botany, School of
General Studies, Australian National University, Box 197. P.O., City, Canberra,
A.C.T., 2601.
Pulley, Mrs. Jean May, B.Sc.Agr., Dip.Ed., 12 Clisby Close, Cook, A.C.T., 2600.
Racek, Albrecht Adalbert, Dr.rernat. (Brno, Czechoslovakia), School of Biological
Sciences, Department of Zoology, Sydney University, 2006.
Rade, Janis, M.Sc., Flat 28a, 601 St. Kilda Road, Melbourne, Victoria, 3000.
Ralph, Professor Bernhard John Frederick, B.Sc., Ph.D. (Liverpool), A.A.C.I., School of
Biological Sciences, University of New South Wales, P.O. Box 1, Kensington, N.S.W.,
2033.
Ramsay, Mrs. Helen Patricia, M.Se., Ph.D., School of Biological Sciences, University of
New South Wales, P.O. Box 1, Kensington, N.S.W., 2033.
Reye, Hric James, M.B., B.S. (Univ. Q’ld.), Entomology Department, University of
Queensland, St. Lucia, Queensland, 4067.
Reynolds, Miss Judith Louise, Ph.D., 118 Homer Street, Earlwood, N.S.W., 2206.
Richards, Miss Aola Mary, M.Sc. (Hons.), Ph.D. (N.Z.), School of Biological Sciences,
University of New South Wales, P.O. Box 1, Kensington, N.S.W., 2033.
Richards, Bryant Neville, B.Se.For., Ph.D. (Q’ld.), 21 College Avenue, Armidale, N.S.W..,
2350.
Richardson, Barry John, 12 Bowden Street, Parramatta North N.S.W., 2150.
Richardson, Laurence Robert, M.Se., Ph.D., F.R.S.N.Z., 4 Bacon Street, Grafton, N.S.W..,
2460.
Riek, Edgar Frederick, B.Sc., Division of Entomology, C.8.I1.R.O., P.O. Box 109, City,
Canberra, A.C.T., 2601. .
Rigby, John Francis, B.Sc., 152 Holland Road, Blackburn South, Victoria, 3130.
*Robertson, Sir Rutherford Ness, C.M.G., F.R.S., B.Sc., Ph.D., F.A.A., Professor of Botany,
University of Adelaide, Adelaide, South Australia, 5000.
Rodd, Anthony Norman, Royal Botanic Gardens, Sydney, 2000.
Rothwell, Albert, D.P.A., 11 Bonnie View Street, Cronulla, N.S.W., 2230.
Salkilld, Barry William, Dip.Soc.Stud. (Univ. Syd.), 71 Beresford Road, Thornleigh,
N.S.W., 2120.
*Salter, Keith Eric Wellesley, B.Sc.. School of Biological Sciences, Department of Zoology,
Sydney University, 2006.
Sands, Miss Valerie Elizabeth, M.Sc., Flat 3, 2 William Street, Dunedin, New Zealand.
*Scammell, George Vance, B.Sc., 7 David Street, Clifton Gardens, N.S.W., 2088..
Selkirk, David Robert, School of Biological Sciences, Botany Building, Sydney University,
2006.
Selwood, Mrs. Lynne, B.Sc., Ph.D., (née Bedford), School of Anatomy, University of New
South Wales, P.O. Box 1, Kensington, N.S.W., 2033.
*Sharp, Kenneth Raeburn, B.Sc., Eng. Geology, 8.M.H.E.A., Cooma, N.S.W., 2630.
Shaw, Miss Dorothy Edith, M.Sc.Agr.. Ph.D., Department of Agriculture, Stock and
Fisheries, Port Moresby, Papua-New Guinea.
Sherrard, Mrs. Kathleen Margaret, M.Sc., 43 Robertson Road, Centennial Park, Sydney,
2021.
Shipp, Erik, Ph.D., 23 Princes Street, Turramurra, N.S.W., 2074.
Simons, John Ronald, M.Sec., Ph.D., 242 Kissing Point Road, Turramurra, N.S.W., 2074.
Slack-Smith, Richard J., 7 Kingston Street, Shenton Park, Western Australia, 6008.
Smith, Eugene Thomas, 22 Talmage Street, Sunshine, Victoria, 3020.
Smith-White, Spencer, D.Sc.Agr., F.A.A., 51 Abingdon Road, Roseville, N.S.W., 2069.
South, Stanley A., B.Sc., 47 Miowera Road, Turramurra, N.S.W., 2074.
Southcott, Ronald Vernon, M.B., B.S., 13 Jasper Street, Hyde Park, South Australia,
5061. :
1937
1960
1968
1932
1956
1965
1962
1965
1952
1962
1962
1965
1940
1950
1950
1956
1960
1949
1944
1968
1943
1946
1921
1965
1952
1949
1917
1930
1940
1934
1961
1952
1909
1967
1946
1947
1911
1966
1936
LIST OF MEMBERS 495
Spencer, Mrs. Dora Margaret, M.Sc. (née Cumpston), No. 1 George Street, Tenterfield,
N.S.W., 2372.
Staff, Ian Allen, B.Se., Dip.Ed., Department of Botany, School of Biological Sciences,
La Trobe University, Bundoora, Victoria, 3083.
Stanbury, Peter John, Ph.D., The Macleay Museum, School of Biological Sciences, Sydney
University, 2006.
Stead, Mrs. Thistle Yolette, B.Sc. (née Harris), 14 Pacific Street, Watson’s Bay, N.S.W.
2030.
Stephenson, Neville George, M.Sc. (N.Z.), Ph.D. (Lond.), School of Biological Sciences,
Department of Zoology, Sydney University, 2006.
Stephenson, . Professor William, B.Sc. (Hons.), Ph.D. (Durham, Eng.), Diploma of
Education, member Aust. Coll. Educ., Fellow Zoological Society, Department of
Zoology, University of Queensland, St. Lucia, Queensland, 4067.
Strahan, Ronald, M.Sc., Director, Taronga Zoo, P.O. Box 20, Mosman; N.S.W., 2088.
Straughan, Mrs. Isdale Margaret, B.Sc., Ph.D., Allan Hancock Foundation, University of
Southern California, University Park, Los Angeles, California, 90007, U.S.A.
Sullivan, George Emmerson, M.Sc. (N.Z.), Ph.D., Department of Histology and
Embryology, Sydney University, 2006.
Sweeney, Anthony William, Malaria Institute, Public Health Department, Rabaul,
Territory: of Papua and New Guinea.
*Swinbourne, Robert Frederick George, 4 Leeds Avenue, Northfield, South Australia, 5085.
Talbot, Frank Hamilton, M.Se., Ph.D., Australian Museum, P.O. Box A285, Sydney
South, N.S.W., 2000.
Taylor, Keith Lind, B.Sc.Agr., c/- C.S.I.R.O., Division of Entomology, Stowell Avenue,
Hobart, Tasmania, 7000.
Tchan, Professor Yao-tseng, Dr., es Sciences (Paris), Department of Microbiology, Sydney
University, 2006.
Thompson, Mrs. Joy Gardiner, B.Sc.Agr. (née Garden), 21 Middle Head Road, Mosman,
N.S.W., 2088.
Thomson, James Miln, D.Sc. (W.A.), Department of Zoology, University of Queensland,
St. Lucia, Queensland, 4067.
Thorne, Alan Gordon, B.A., Department of Anatomy, Sydney University, 2006.
Thorp, Mrs. Dorothy Aubourne, B.Sc. (Lond.), Ph.D., ‘‘ Sylvan Close’’, Mt. Wilson,
N.S.W., 2740.
Thorpe, Ellis William Ray, B.Sc., University of New England, Armidale, N.S.W., 2350.
Timms, Brian Victor, B.Sc.(Hons.), Zoology Department, Monash University, P.O. Box 92,
Clayton, Victoria, 3168.
Tindale, Miss Mary Douglas, D.Sc., 60 Spruson Street, Neutral Bay, N.S.W., 2089.
Tipper, John Duncan, A.M.I.E.Aust., 26 Kuring-gai Avenue, Turramurra, N.S.W., 2074.
*Troughton, Ellis Le Geyt, C.M.Z.S., F.R.Z.S., c/- Australian Museum, P.O. Box A285,
Sydney South, N.S.W., 2000.
Tucker, Richard, B.V.Sc., Dr.Vet.M., Veterinary School, Department of Veterinary
Anatomy, University of Queensland, St. Lucia, Queensland, 4067.
Valder, Peter George, B.Sc.Agr., Ph.D. (Camb.), School of Biological Sciences, Depart-
ment of Botany, Sydney University, 2006.
Vallance, Professor Thomas George, B.Sc., Ph.D., Department of Geology and Geophysics,
Sydney University, 2006.
Veitch, Robert, B.Sc., F.R.E.S., 24 Sefton Avenue, Clayfield, Queensland, 4011.
Vickery, Miss Joyce Winifred, M.B.E., D.Sc., Royal Botanic Gardens, Sydney, 2000.
Vincent, Professor James Matthew, D.Sc.Agr., Dip.Bact., Department of Microbiology,
School of Biological Sciences, University of New South Wales, P.O. Box 1, Kensington,
N.S.W., 2033.
*Voisey, Professor Alan Heywceod, D.Sec., School of Earth Sciences, Macquarie University,
North Ryde, N.S.W., 2113.
Walker, Donald, B.Sc., M.A., Ph.D., F.L.S., 18 Cobby Street, Campbell, Canberra, A.C.T.,
2601.
Walker, John, B.Sc.Agr., Biological Branch, N.S.W. Department of Agriculture, Private
Mail Bag No. 10, Rydalmere, N.S.W., 2116.
Walkom, Arthur Bache, D.Sc., 5/521 Pacific Highway, Killara, N.S.W., 2071.
Wallace, Michael Joseph, B.Sc., 66 Parry Avenue, Beverly Hills, N.S.W., 2209.
Wallace, Murray McCadam Hay, B.Sc., C.S.I.R.O., W.A. Regional Laboratory, Nedlands,
Western Australia, 6009.
Ward, Mrs. Judith, B.Sc., 16 Mortimer Avenue, New Town, Hobart, Tasmania, 7008.
Wardlaw, Henry Sloane Halcro, D.Sc., F.R.A.C.I., 71 McIntosh Street, Gordon, N.S.W.,
2072.
Wass, Robin Edgar, B.Sc. (Hons., Q’ld.), Ph.D., Department of Geology and Geophysics,
Sydney University, 2006.
Waterhouse, Douglas Frew, D.Sc., F.R.S., C.S.I.R.O., P.O. Box 109, City, Canberra, A.C.T.,
2601.
496
1947,
1927
1941
1964
1963
1963
1946
1963
1926
1962
1960
1954
1954
1960
1952
1950
1947
1968
1965
1964
1964
1965
1949
LIST OF MEMBERS
*Waterhouse, John Teast, B.Sc., School of Biological Sciences, University of New South
Wales, P.O. Box 1, Kensington, N.S.W., 2033.
Waterhouse, Professor Walter Lawry, C.M.G., D.Se.Agr., M.C., D.I.C., F.L.S., F.A.A.,
30 Chelmsford Avenue, Lindfield, N.S.W.., 2070.
Watson, Professor Irvine Armstrong, Ph.D., B.Sc.Agr., Faculty of Agriculture, Sydney
University, 2006.
Webb, Mrs. Marie Valma, B.Sc., 30 Talfourd Street, Glebe, N.S.W., 2037.
Webby, Barry Deane, Ph.D., M.Sc., Department of Geology and Geophysics, Sydney
University, 2006.
Webster, Miss Elsie May, 20 Tiley Street, Cammeray, N.S.W., 2062.
Wharton, Ronald Harry, M.Sc., Ph.D., C.S.I.R.O., 677 Fairfield Road, Yeerongpilly,
Queensland, 4105.
White, Mrs. Mary Elizabeth, M.Sc., 7 Ferry Street, Hunter’s Hill, N.S.W., 2110.
*Whitley, Gilbert Percy, F.R.Z.S., Australian Museum, P.O. Box A285, Sydney South,
: N.S.W., 2000.
Whitten, Maxwell John, Ph.D., C.S.I.R.O., Division of Entomology, P.O. Box 109, City,
Canberra, A.C.T., 2601.
Wildon, David Conrad, B.Sc.Agr., Box 108, Rozelle, N.S.W., 2039.
Williams John Beaumont, B.Sc., University of New England, Armidale, N.S.W., 2350.
Williams, Mrs. Mary Beth, B.Sc. (née Macdonald), 902D Rockvale Road, Armidale, N.S.W.,
2350.
Williams, Neville John, B.Sc., Zoologisch Laboratorium der Rijksuniversiteit te Groningen,
Rijksstraatweg 78, Haren (Gr.), Nederland.
Williams, Owen Benson, M.Agr.Sc. (Melbourne), c/- C.S.I.R.O., The Ian Clunies Ross
Animal Research Laboratory, P.O. Box 144, Parramatta, NS. W., 2150.
Willis, Jack Lehane, M.Se., A.A.C.1., 26 Inverallan Avenue, Pymble, N.S.W., 2073.
Winkworth, Robert Ernest, Rangelands Research Unit, C.S.I.R.O. ,P.O. Box 109, Canberra
City, A.C.T., 2601.
Wood, Alec Edward, B.Sc.Agr., Ph.D., 25 George Street, Bexley, N.S.W., 2207.
Woodward, Thomas Emmanuel, M.Sc. (N.Z.), Ph.D. (Lond.), D.1I.C., Department of
Entomology, University of Queensland, St. Lucia, Queensland, 4067.
Wright, Anthony James Taperell, B.Sc., Geology Department, Victoria University of
Wellington, P.O. Box 196, Wellington, C.1., New Zealand.
Yaldwyn, John Cameron, Ph.D.(N.Z.), M.Sc., Dominion Museum, Private Bag,
Wellington, New Zealand. :
Young, Graham Rhys, 8 Spark Street, Harlwood, N.S.W., 2206.
CORRESPONDING MEMBER
Jensen, Hans Laurits, D.Sc.Agr. (Copenhagen), State Laboratory of Plant Culture,
Department of Bacteriology, Lyngby, Denmark.
497
LIST OF PLATES
PROCEEDINGS, 1968
I—Fig. 1. Miwvophyes fasciolatus; Fig. 2. Mixophyes iteratus.
Iil—Fig. 1. Mivophyes balbus; Fig. 2. Mixophyes schevilli.
Il1I-V.—The mucosa of the stomach of the wombat.
VI—An enclosure on a gilgai on the Burt Plain, 19th April, 1961.
VII.—An enclosure on a gilgai on the Burt Plain, 27th April, 1961.
VIII.—Pattern of hairs on leaves of Hragrostis species.
IX—XI.—The Lower and Middle Palaeozoic stratigraphy and Sedimentary
Tectonics of the Sofala—Hill End—Euchareena region, N.S.W.
XII.—Geological Map of the Euchareena—Hill End-Sofala area.
XITI.—Aphrophyllum hallense Smith and A. smithi, sp. nov.
XIV-XV.—Permian faunas and sediments from the South Marulan district,
New South Wales.
XVI.—Chimaerism for mildew reaction type in wheat.
XVIT.—Phenotypic expression of pubescent glumes in wheat.
XVITI-XX.—The vegetation of the Boorabbin and Lake Johnston areas.
XXI.—Boorabbin—Vegetation survey of Western Australia.
XXII.—Lake Johnston—Vegetation survey of Western Australia.
XXITTI-XXVITI—Smuts of Hchinochloa spp.
XXIX.—Patiriella vivipara, sp. nov.
XX X-XXXI.—The nasal mites of Queensland birds.
LIST OF NEW GENERA AND NEW SPECIES
VoL. 93
New Genera
Page
Bdellasimilis (Planariidae) si af ye a: nts a2 ee Ol
Pallidotettia (Rhaphidophoridae) bis Jee rs Sy Hs oe AG
New Species
Page Page
accipitris (Ophthalmognathus) 390 myzomelae (Boydaia) .. .. 383
artami (Ruandanyssus) .. .. 330 neochmiae (Ptilonyssus) rome S41
balbus (Mixophyes) Hick Sere reas, neosittae (Sternostoma) co BCC
barwicki (Bdellasimilis) one OL nullarborensis (Pallidotettir) 48
corcoracis (Ptilonyssus) .. 345 orthonychus (Ptilonyssus) .. 354
gerygonae (Ptilonyssus) Stal OOS prima (Hlmmata) .. .. .. . 220
iteratus (Mixophyes) .. ~.. 54 setosae (Ptilonyssus) .. .. 362
maluri (Boydaia) .. .. .. 3886 smithi (Aphrophyllum) .. .. 198
megaloprepiae (Tinaminyssus) 316 struthideae (Ptilonyssus) .. 348
monarchae (Ptilonyssus) .. 360 vivipara (Patiriella) _ ae OA:
myristicivorae (Tinaminyssus) 316 welchi (Tinaminyssus) .. .. 319
498
Page Page
Abstract of Proceedings . 482 Dartnall, A. J., Pawson, D. L., Pope,
Agnesiidae Huntsman 1912, a review
of the family; with particular
reference to Agnesia omer
Michaelsen 1898
Anderson, D. T., and Rossiter, G. T.
Hatching and larval development
of Dissonus nudiventris Kabata
(Copepoda, Fam. Dissonidae), a
gill parasite of the Port Jackson
shark, 476—Hatching and larval
development of Haplostomella
australiensis Gotto (Copepoda,
Fam. Ascidicolidae), a parasite
of the ascidian aa Giese
Herdman
Annual General nreetine E
Aphrophyllum (Rugosa), from Lower
Carboniferous limestones near
Bingara, N.S.W. AE
Australian decapod Grustacce: the
constitution, distribution and
relationships of the
Baker, E. P., see McIntosh, R. A. and
Baker, of P.
Balance Sheets for the year ending
29th February, 1968 a
Beard, J. S., The vegetation of the
Boorabbin and Lake Johnston
areas, Western Australia
Birds, Queensland, the nasal mites of
Burbidge, Nancy T., Notes on Vitta-
dinia triloba sens. lat. (Com-
positae)
Chippendale, G. M., The plants, erared
by red kangaroos,. Megaleia rufa
(Desmarest) in Central Aust-
ralia ‘ a he Hs
Chromosome location and linkage
studies involving the Pm3 locus
for powdery mildew resistance
in wheat one Shs See
Climacograptus bicornis with a
modified basal assemblage, on the
first occurrence of a, in Australia
Collie, Rev. R, ee fg of,
repaired
Congratulations
Constitution, Gictripacion ana as
tionships of the Australian
decapod Crustacea BiG j
Dandie, Alison K., reappointed
Linnean Macleay Fellow in
Botany for 1969
Dartnall, A. J., A viviparous species
of Patiriella (Asteroidea, Aster-
inidae) from Tasmania ..
. 444
164
5-7
. 239
297
437
98
. 164
487
. 294
Elizabeth C., and Smith, B. J.,
Replacement name for the pre-
occupied genus name Odinia
Perrier 1885 on Be railat
Davis, Gwenda L., The emi yeloee of
EHpaltes australis Less. (Com-
positae) ay Ay 4 .. 184
Dissonus nudiventris Kabata, hatch-
ing and larval development of, a
gill ina of the Port Jackson
shark ; : . 476
Domrow, R., The nasal mites of
Queensland birds (Acari: Der-
manyssidae, Hreynetidae and
Epidermoptidae) cf .. 297
Echinochloa spp., a study of some
smuts of 3 ae as B95 hell
Elections . 8, 482-483, 485-487
Epaltes australis Less., the oe
ology of ih 5 .. 184
Exhibits—see Notes and Exhibits.
Frith, H. J., Sir William Macleay
Memorial Lecture, 1968. Wildlife
Conservation a .. 270
Fullerton, R. A., and epee
R. F. N., A study of some smuts
of Echinochloa spp. Bs .. 281
George, HE. P., see Milton, G. W.,
Hingson, D. J., and George, E. P.
Goldman, Judy, Hill, L., and Stan-
bury, P. J., Type specimens in
the Macleay Museum, University
of Sydney. II. pes and
Reptiles 55 .. 427
Gould, I. G., see Wass, R. E., and
Gould, I. G.
Griffin, D. J. G., and Yaldwyn, J. C.,
The constitution, distribution and
relationships of the Australian
decapod Crustacea ie .. 164
Halocynthia Verrill 1879, a review
of the genus Ba: Ne apes HO
Haplostomella australiensis Gotto
(Copepoda, Fam. Ascidicolidae),
hatching and larval development
of, a parasite of the ascidian
Styela etheridgii Herdman. .. 464
Hatching and larval development of
Dissonus nudiventris Kabata
(Copepoda, Fam. Dissonidae), a
gill nee ak of the Port Jackson
shark ¢ : d .. 476
INDHX
Page
Hatching and larval development
of Haplostomella australiensis
Gotto (Copepoda, Fam. Ascidi-
colidae), a parasite of the
ascidian a Eno Herd-
man .. .. 464
Hill, L., see coe ae Hill,
L., and Stanbury, P. J.
Hingson, D. J., see Milton, G. W.,
Hingson, D. J., and George, E.P.
Hingson, D. J., and Milton, G.W.,
- The mucosa of the stomach of
the wombat (Vombatus hirsutus)
with special reference to the
cardiogastric gland re ; 69
Hotchkiss, A. T., see Notes a
Exhibits.
Johnson, L. A. S§S., Presidential
Address. Rainbow’s End: .the
quest for an optimal taxonomy 8
Jull, R. K. Aphrophyllum (Rugosa)
from Lower Carboniferous lime-
stones near Bingara, N.S.W. .. 193
Kangaroos, red, Megaleia rufa (Des-
marest), the plants grazed HP
in Central Australia er 98
Kott, Patricia, A review of the aie
Agnesiidae Huntsman 1912; with
particular reference to Agnesia
glaciata Michaelsen 1898, 444—
A review of the genus dHalo-
cynthia Verrill 1879 oe been LO
Langdon, R. F. N., see Fullarton,
R. A., and Langdon R. F. N.
Lecturettes . 1, 484-487
Library Accessions . 1, 482-487
Linnean Macleay Fellowship:
Reappointment of Miss Alison K.
Dandie for 1968, 2—summary
of work, 2—applications in-
vited for 1969, 486—487—
reappointment for 1969 .. 487
Linnean Macleay Lectureship in
Microbiology:
Report of work by Dr. Y. T.
Tchan for year ending 31st
December, 1967
Lower and Middle Palaeozic yee
graphy and sedimentary tec-
tonics of the Sofala-Hill End-
Huchareena region, N.S.W. aa Jt
Macleay Museum, University of
Sydney, type specimens in the,
I. Fishes, 203—II. Amphibia and
Reptiles, 427—III, Birds, 457—
IV. Mammals we ee e462,
McIntosh, R. A., and Baker, E. P.,
Chromosome location and linkage
studies involving the Pm3 locus
for powdery mildew resistance in
wheat ie aA if 50 LB
Members, List of Betis mene .. 489
Milton, G. W., see Hingson, D. J.,
and Milton, G. W.
- Nasal
499
Page
Milton, G. W., Hingson, ID) dey, hava!
George, BE. P., The _ secretory
capacity of the stomach of the
wombat (Vombatus hirsutus)
and the cardiogastric gland
Mites, nasal, of Queensland birds ..
Mixophyes, a taxonomic review of
the genus
Moors, H., On the first occurrence of
a Climacograptus bicornis with a
modified basal aBESTIBIGES: in
- Australia ;
Mucosa of the stomach of the ree
(Vombatus hirsutus), with
special reference to the cardio-
gastric gland
mites of een eae
(Acari: Dermanyssidae, Erey-
netidae and Epidermoptidae)
New bdellourid-like triclad turbellar-
jan ectoconsortic on Murray
River Chelonia aa 3
Notes and Exhibits:
Hotchkiss, A. T.—Discussion of
the ripening and after ripening
in fruits of Ruppia ne
Dumort
Newman, I. 7 —Ohserna tions re
laying of a sewer line beside
the bed of the Lane Cove
River near its headwaters
between Wahroonga and
Normanhurst which means
great scarring and destruc-
tion
Vallance, T. Go Reference to fhe
centenary (1967) of the Rey.
W. B. Clarke’s work
“Remarks on the Sedimen-
tary Formations of New
South Wales”
Whitley, G. P.—Exhibition Anil
comments upon some Japan-
ese papers on poisonous
crabs, 488—Exhibition of two
unpublished photographic
portraits, from the Aust-
ralian Museum’s_ archives,
of George French Angas and
Gerard Krefft .. oe
Notes on Vittadenia triloba sens.
lat. (Compositae)
Obituary Notice:
Professor P. D. F. Murray
On the first occurrence of a
Climacograptus bicornis with a
modified basal
Australia
Packham, G. H., The Lower and
Middle Paleozoic stratigraphy
and sedimentary tectonics of
the Sofala-Hill Hnd-Huchareena
region, N.S.W. 5
Patiriella from Tasmania,
ay Vilva=
parous species of ..
60
297
52
227
69
5 ABT
90
488
483
488
ay sree in
» |
Iii
. 294
500
INDEX
Page
Pawson, D. L., see Dartnall, A. J.,
Pawson, D. L., Pope, Elizabeth C.,
and Smith, B. J.
Peacock, W. J., reference to award
of Edgeworth David Medal by
the Royal Society of N.S.W.,
conjointly with Dr. D. M. Green ..
Permian faunas and sediments from
the South Marulan district,
N.S.W. Pe a ;
Plants grazed by red kangaroos,
Megaleia rufa (Desmarest), in
‘Central Australia Ly Bele
Plates, List of ..
Pope, Elizabeth C., see Dartnall, A. J.,
Pawson, D. ie Pope, Elizabeth
C., and Smith, B. J. ..
Port Jackson shark, hatching ana
larval development of Dissonus
nudiventris Kabata, a gill
parasite of the ne sat
Presidential Address , 3
Raggatt, Sir H., reference to death .
Rainbow’s End: the quest for an
optimal taxonomy (Presidential
Address)
Reference to deaths
Replacement name for the _ pre-
occupied genus name Odinia
Perrier 1885
Report on the Affairs cof the Society
Review of the family Agnesiidae
Huntsman 1912; with particular
reference to Agnesia beac
Michaelsen 1898 2 nit
Review of the genus
Verrill 1879
Rhaphidophoridae (Oren of
Australia. 7. Pallidotettiz, a new
genus from the Nullarbor Plain,
South-western Australia
Richards, Aola M., The Rhaphido-
phoridae pole aged of Aust-
ralia. 7
Richardson, L. R., A new pasitouria:
like triclad turbellarian ectocon-
sortic on Murray River Chelonia
Rossiter, G. T., see Anderson, D. T.,
and Rossiter, G. T.
Rules of Society, revised and Te-
printed
Science House
Secretory capacity of Ne Shes pth of
the wombat (Vombatus hirsutus )
and the cardiogastric gland
Sir William Macleay Memorial
Lecture, 1968. Wildlife Conserva-
tion £
Jal ctocynthia
98
5 BOT
5 eqilal
. 444
76
46
46
90
. 270
Page
Smith, B. J., see Dartnall, A. J.,
Pawson, D. L., Pope, Elizabeth C.,
and Smith, B. J.
Smuts of Hchinochloa spp., a paper’
of some
Stanbury, P. J., Tape STOR OTS in
the Macleay Museum, University
of Sydney, I. Fishes, 203—IITI.
Birds, 457—IV. Mammals, 462—
see Goldman, Judy, Hill, L., and
Stanbury, P. J.
Straughan, I. R. A taxonomic review
of the genus Mixophyes (Anura,
Leptodactylidae)
Study of some smuts of eTneniee
Spp.
Styela etheridgii Herdman, hatching
and larval development of
Haplostomella australiensis Gotto
(Copepoda, Fam. Ascidicolidae),
a parasite of the ascidian
Talbot, F. H., elected a member of
Council es iG. ;
Taxonomic review of the genus
Mixophyes nee: Leptodacty-
lidae )
Turbellarian, a new adellonnine like
triclad, ectoconsortic on Murray
River Chelonia FS tc
Type specimens in the Macleay
Museum, University of Sydney,
I. Fishes, 203—II. Amphibians
and Reptiles, 427—III. Birds,
457—IV. Mammals
Vallance, T. G., elected Preedenn
3—see Notes and Exhibits.
Vegetation of the Boorabbin and
Lake Johnston areas, Western
Australia
Vittadinia triloba sens. Se. Nore on
Viviparous species of Patiriella
(Asteroidea, Asterinidae) from
Tasmania 2
Wass, R. E., and Gould, th G., Denenian
faunas “and sediments ‘from the
South Marulan district, N.S.W.
Wheat, chromosome location and
linkage studies involving the
Pm3 locus for powdery mildew
resistance in : ot A
Whitley, G. P., see Notes and
Exhibits.
Wildlife Conservation. Sir William
Macleay Memorial Lecture, 1968
Wombat (Vombatus hirsutus), the
mucosa of the stomach of the,
with special reference to the
cardiogastric gland As
Wombat (Vombatus hirsutus), the
secretory capacity of the stomach
of, and the ecardiogastric gland
Yaldwyn, J. C., see Griffin, D. J. G.,
and Yaldwyn, J. C.
281
52
281
. 464
. 485
52
90
. 462
. 239
439
. 294
212
. 232
270
69
60
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